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A camouflaged diversity: taxonomic revision of the thorny lacewing subfamily Symphrasinae (Neuroptera, Rhachiberothidae)
expand article infoAdrian Ardila-Camacho§, Renato José Pires Machado|, Michael Ohl, Atilano Contreras-Ramos
‡ Instituto de Biología, Ciudad de México, Mexico
§ Universidad Distrital “Francisco José de Caldas”, Bogotá, Colombia
| Universidade Federal do Paraná, Curitiba, Brazil
¶ Museum für Naturkunde Berlin, Berlin, Germany
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Abstract

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.

Key words

Lacewings, Mantispoidea, morphology, New World, systematics

Dedication

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.

Introduction

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 (Engel et al. 2018; Ardila-Camacho et al. 2021a). Understanding of the phylogenetic relationships between these families remain contentious, in part because of a highly diverse morphology, disparate distribution, relatively low diversity, and a long evolutionary history with a considerable number of extinct lineages (Snyman et al. 2020; Lu et al. 2020; Ardila-Camacho et al. 2021a). The traditional scheme of classification of Mantispidae considers four subfamilies, namely Symphrasinae, Drepanicinae, Calomantispinae, and Mantispidae, of which the first has been considered as sister of the latter three for long time and often referred to as “the most primitive” subfamily of mantidflies (Penny and da Costa 1983; Lambkin 1986a, b; Liu et al. 2015; Shi et al. 2019). Moreover, the Rhachiberothidae, which was originally proposed as a subfamily of Berothidae, includes a single extant subfamily, i.e., the Rhachiberothinae, plus the extinct Paraberothinae from the Cretaceous. This group has been also considered as a subfamily of Mantispidae, sister to the rest of mantispid subfamilies (Willmann 1990), or a separated family, either sister to Berothidae (Aspöck and Mansell 1994; Song et al. 2019) or sister to Mantispidae (Liu et al. 2015; Wang et al. 2017).

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 (Winterton et al. 2018). In connection with these results, consistent morphological support for a new concept of Rhachiberothidae, including Symphrasinae was proposed by Ardila-Camacho et al. (2021a). This study recovered the Rhachiberothidae as a monophyletic group with Symphrasinae sister to the extinct Paraberothinae in a clade, which is sister to the Rhachiberothinae based on the presence of modified setae on the closing surface of the forebasitarsus and the presence of forebasitarsal Stitz organ. Also, this family was recovered as sister to the true Mantispidae (i.e., Mantispinae + (Calomantispinae + Drepanicinae)) whose monophyly is strongly supported by many characters of the raptorial apparatus, wing venation, and genitalia. The hypothesis of the sister-group relationship between these families assumes a single origin of the raptorial condition within the superfamily, and the monophyly of each subfamily and their relationships itself are supported by characters of the raptorial apparatus, primarily of the foretarsus (Ardila-Camacho et al. 2021a).

The subfamily Symphrasinae was proposed by Lambkin (1986a), although it was originally created by Navás (1909) as a tribe of Mantispidae, i.e., the Symphrasini. Since then, this group has undergone numerous nomenclatural changes due to a considerable homonymy and synonymy of its genera (Penny 1982b). Currently this group includes three extant genera restricted to the New World, namely, Anchieta Navás, 1909 with, previous to the current revision, eight valid species ranging from Panama to southern Brazil, Plega Navás, 1928 composed of 17 valid species distributed from southwestern United States to southern Brazil, and Trichoscelia Westwood, 1852 with 17 valid species known from southern Mexico to Argentina (Ardila-Camacho et al. 2018; Machado 2018; Ardila-Camacho et al. 2019). Nevertheless, the subfamily has a relatively rich fossil record, with six genera, i.e., Archaeosymphrasis Shi et al. 2019, Habrosymphrasis Shi et al. 2019, Parasymphrasites Lu et al. 2020, Haplosymphrasites Lu et al. 2020, Parvosymphrasites Li et al. 2023 and Proplega Li et al. 2023 from the Upper Cretaceous deposits of Myanmar (Shi et al. 2019; Lu et al. 2020; Li et al. 2023). Among these, the first three were proposed as stem-group representatives of the Symphrasinae (Shi et al. 2019; Lu et al. 2020). On the other hand, Haplosymphrasites and Symphrasites Wedmann & Makarkin, 2007 from the Middle Eocene of Germany are considered the crown-group of Symphrasinae together the extant genera (Wedmann and Makarkin 2007; Shi et al. 2019; Lu et al. 2020).

The type genus of this subfamily, i.e., Symphrasis was proposed by Hagen (1877) to accommodate two species: S. signata and S. myrapetrella (Westwood, 1867), although according to Enderlein (1910), the type species of the genus is actually Raphidia varia Walker, 1853. The genus Plega was synonymized with Symphrasis by Tjeder (1959), although Parker and Stange (1965) proposed Symphrasis as a synonym of Trichoscelia (Penny 1982b) and reestablished Plega after finding consistent differences between both genera. Moreover, Trichoscelia and Symphrasis were synonymized with Anisoptera Schneider by Gerstaecker (1888), with the synonymy of the former genus confirmed by Banks (1913). Nonetheless, Anisoptera is not an available name, and its type species belongs to Anchieta. The latter genus was synonymized with Anisoptera by Enderlein (1910), and this synonymy was confirmed by Banks (1913) and Penny (1982b). Meanwhile, the generic names Platymantispa Rehn, 1939 and Anisopterana Strand, 1942 were unnecessarily proposed and independently created, because Anisoptera is a junior homonym. Considering this background of confusing nomenclatural changes, the oldest available name for the subfamily, i.e., Symphrasini prevailed, because according to the article 40 of the International Code of Zoological Nomenclature, the invalidity of its type genus was not recorded before 1961 (Parker and Stange 1965; Lambkin 1986a).

The phylogenetic relationships of the genera in the Symphrasinae are not well understood, and the boundaries of the genera are not clearly outlined (Liu et al. 2015). The first hypothesis of relationships between the symphrasine genera recovered Trichoscelia sister to Anchieta + Plega (Penny and da Costa 1983), which was also recovered by Willmann (1990) and Liu et al. (2015) based on morphology and total evidence analyses, respectively. Further morphological studies including extant and extinct genera recovered Archaeosymphrasis, Habrosymphrasis and Symphrasites as transitional genera between Mesomantispinae and the extant crown-group of Symphrasinae, where Trichoscelia was recovered as sister of the other two genera (Shi et al. 2019). In a subsequent research, Shi et al. (2020) presented hypotheses from different morphological datasets, mostly without clear relationships between extinct and extant genera, and between extant genera. Such work recovered Archaeosymphrasis and Habrosymphrasis in a clade sister to Plega and a trichotomy of Symphrasites, Anchieta, and Trichoscelia from a matrix, where the genitalia characters were excluded. Additionally, from the same dataset excluding genitalia characters and poorly preserved fossils, Plega was recovered as sister to Anchieta + Trichoscelia (Shi et al. 2020). Nearly simultaneously, Lu et al. (2020), recovered Parasymphrasites and Habrosymphrasis as stem group Symphrasinae, and Symphrasites as sister of a polytomy between the extant genera and Haplosymphrasites.

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 (Rehn 1939b; Penny 1982a; Penny and da Costa 1983). Nevertheless, morphological descriptions for many species lack enough details and the taxonomic status of the majority of them needs to be clarified. Consequently, a reliable taxonomic identification of the species is often difficult to achieve, particularly in northern South America, Mexico and Central America, where the group has its higher diversity. For these reasons, the objectives of this study are to test the monophyly of the extant genera in the Symphrasinae and to infer their phylogenetic relationships based on morphological data. Additionally, we reconstruct the phylogenetic relationships between the species in the Symphrasinae and provide diagnosis and descriptions for all of them, clarifying their taxonomic status.

Materials and methods

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 (Ardila-Camacho et al. 2019). Genital complexes were placed in a setup composed of a petri dish with a layer of paraffin at the bottom and filled with alcohol. The genitalia were situated at different views in a canal made with the tip of entomological forceps. Genital structures were placed in microvials filled with glycerin for permanent storage. External morphology was studied in a Zeiss Discovery V8 stereomicroscope. High resolution photographs were taken with a ZEISS AxioZoom V16 microscope, and then stacked using the ZENpro201 software. Images were processed using Adobe Photoshop CS6. Morphological terminology and homology followed Ardila-Camacho et al. (2021a), the terms for foreleg morphology are summarized in Fig. 1, and for wing venation and male and female genitalia are depicted in Fig. 2.

Figure 1. 

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.

Figure 2. 

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.

Museums and collections cited across the manuscript

AMNH American Museum of Natural History, New York, USA.

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA

CAS California Academy of Sciences, San Francisco, USA.

CAUD Colección de Artrópodos, Universidad Distrital Francisco José de Caldas, Bogotá, Colombia.

CN Navás collection housed at (MZBS) Museo Zoologia, Barcelona, Spain.

CNIN Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México, Ciudad de México, México.

CSCA California State collection of Arthropods, California Department of Food and Agriculture, Sacramento, California, USA.

CZMA Coleção Zoológica do Maranhão, Caixias, Maranhão, Brazil.

DZUP Coleção Entomológica Padre Jesus Santiago Moure, Curitiba, Paraná, Brazil.

FSCA Florida State Collection of Arthropods, Gainesville, Florida, USA.

IAvH Instituto Alexander von Humbolt, Villa de Leyva, Colombia.

ICN Instituto Nacional de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia.

INPA Coleção Sistematica da Entomologia, Manaus, Amazonas, Brazil.

LEUA Colección de Entomología del Laboratorio de Entomología, Universidad de la Amazonia, Florencia, Caquetá, Colombia.

MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA.

MEFLG Museo Entomológico “Francisco Luis Gallego”, Universidad Nacional de Colombia, Medellín, Colombia.

MHNL Musée des Confluences, Lyon, France.

MHN-UPN Museo de Historia Natural, Universidad Pedagógica Nacional, Bogotá, Colombia.

MNCR Museo Nacional de Costa Rica, Santo Domingo de Heredia, Costa Rica.

MNHN Museum national d’Histoire naturelle, Paris, France.

MPEG Museu Paraense Emilio Goeldi, Belem, Para, Brazil.

MPUJ Museo Javeriano de Ciencias Naturales, Bogotá, Colombia.

MUSENUV Museo Entomológico de la Universidad del Valle, Cali, Colombia.

MZPW Museum of the Institute of Zoology, Polish Academy of Sciences, Warszawa, Poland.

NHMUK The Natural History Museum, London, United Kingdom.

NHRS Natuhistoriska Riksmuseet, Stockholm, Sweden.

NMBS Naturhistorisches Museum, Bern, Switzerland.

NMNH National Museum of Natural History, Smithsonian Institution, Washington DC, USA.

OUMNH Oxford University Museum of Natural History, Oxford, Great Britain.

SDEI Senckenberg Deutsches Entomologisches Institut, Eberswalde, Germany (formerly in Berlin-Dahlem).

TAMUIC Insect collection of Texas A&M University, Texas, USA.

UAAM University of Arkansas Arthropod Museum, Fayetteville, Arkansas, USA.

UCD Bohart Museum of Entomology, University of California, Davis, California, USA.

UFMG Universidade Federal de Minas Gerais, Belo Horizonte, Brazil.

UNAB Museo de la Facultad de Agronomía Universidad Nacional de Colombia, Bogotá, Colombia.

UMSP Entomology collection of the University of Minnesota, St. Paul, Minnesota, USA.

ZIN Zoological Museum, Academy of Sciences, St. Petersburg, Russia.

ZIMG Ernst-Moritz-Arndt-Universität, Greifswald, Germany.

ZMB Museum für Naturkunde, Berlin, Germany.

Phylogenetic analysis

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 Ardila-Camacho et al. (2021a). The definition of the characters and character states was based on Strong and Lipscomb (1999). A matrix with 102 adult characters and 56 terminals was handled using the software WinClada® (Nixon 2002) and exported as a nexus file to perform the phylogenetic analysis under the method of parsimony with the program TNT® 1.5 (Goloboff and Catalano 2016). The morphological matrix was analyzed using New Technology search set to 50 employing algorithms of sectorial search, ratchet, drift, and tree fusing, finding the minimum tree 40 times. The characters were treated as unordered and with equal weight. For comparison purposes, an analysis of traditional search of TBR (Tree Bisection and Reconnection) based on 100 random addition sequences, plus an analysis of New Technology with implied weighting with a value of the constant k of 10.717 calculated by means of the TNT script “setk.run” (Goloboff et al. 2008) were conducted. Strict consensus of the most parsimonious trees, as well as optimization and analysis of the evolution of the characters were made using WinClada.

The Bremer absolute (Bremer 1994) was used to calculate support of the branches. This index calculates the number of extra steps needed for a consensus branch to collapse. The Bremer support were calculated using suboptimal trees of 1–10 steps with TBR on the software TNT.

The final topology, including all characters, states and Bremer values, was edited using Adobe Illustrator CS® software.

Morphological characters employed to infer the phylogenetic relationships of the genera and species of Symphrasinae

Head

  1. Ratio of height of vertexal region above compound eyes to distance between upper limit of compound eyes and frontoclypeal ridge : (0) 1:1; (1) 0.5:1; (2) 0.3:1 or 0.2:1.
  2. Overall shape of head, frontal view : (0) globular; (1) ovoid; (2) quadrangular; (3) diamond-shaped.
  3. Shape of frontal sclerite (frons), frontal view : (0) subquadrate; (1) pentagonal; (2) rectangular; (3) bar-shaped.
  4. Ratio of the compound eye width to interocular distance at toruli level, frontal view : (0) ~ 0.75:1; (1) 0.5:1; (2) <0.5:1.
  5. Ocular plane morphology : (0) anteriorly curved inwards; (1) concave; (2) straight.
  6. Antennal scape : (0) as long as wide; (1) 1.5 × as long as wide; (2) ≥2.0× as long as wide.
  7. Basal flagellomeres : (0) as wide as long; (1) 1.5 × as wide as long; (2) two to three × as wide as long; (3) >3 × as wide as long (discoidal).
  8. Shape of palpimacula : (0) fusiform; (1) sulcate; (2) broadly ovoid.
  9. Hypostomal bridge : (0) absent; (1) present.
  10. Supraantenal region : (0) unmodified; (1) with fused tubercles; (2) with lateral, setose protuberances.
  11. Coronal suture : (0) conspicuous; (1) indistinct.

Prothorax

  1. Procoxal insertion : (0) at anterior apex; (1) at posterior end; (2) approximately at mid-length.
  2. Prothorax : (0) two × as long as wide; (1) approximately as long as wide; (2) 1.5 × as long as wide.
  3. Precoxal bridge : (0) present; (1) absent. Uninformative
  4. Pronotum structure : (0) tubular (i.e., posteroventrally fused); (1) shield-shaped.
  5. Ornamentation of the pronotum : (0) coarsely wrinkled; (1) with posterior, transversal furrow; (2) with three uneven, raised areas; (3) with short, raised posterior area; (4) with lateral, short protuberances.
  6. Maculae : (0) present; (1) absent.
  7. Postfurcasternum, shape : (0) narrow-shaped; (1) semi-triangular; (2) plate-shaped; (3) collar-like (i.e., ventrally fused).

Foreleg

  1. Foretrochanter, anterior surface ornamentation : (0) absent; (1) with blunt process.
  2. Trochanter-femur complex : (0) present; (1) absent.
  3. Forefemur, shape : (0) fusiform; (1) subcylindrical (i.e., with approximately subparallel margins); (2) robust, with convex anterior and posterior surfaces.
  4. Posteroventral row of integumentary specializations : (0) not raised; (1) carinate on distal 1/3; (2) carinate on distal ½.
  5. Anteroventral row of integumentary specializations : (0) mostly reduced to apex; (1) fully developed; (2) reduced to proximal region and apex.
  6. Proximal-most process morphology of anteroventral processes row : (0) as a prominent major process; (1) thorn-shaped; (2) as a primary, curved process; (3) not differentiated from other processes.
  7. Posteroventral processes row, arrangement : (0) with two primary processes on distal ½; (1) uniform, having processes with similar shape and size; (2) with single sub-basal, primary process; (3) with two primary processes on proximal ½.
  8. Type of specializations predominant on the rows : (0) spine-shaped; (1) tubercle-shaped.
  9. Stinger-shaped setae on the rows of integumentary specializations : (0) absent; (1) present.
  10. Thickened setae with globular base, adjacent to processes rows : (0) absent; (1) present.
  11. Extension of the posteroventral row of thickened setae with globular base (when present) : (0) present on nearly the entire femoral, ventral surface; (1) present on distal ½ to apex of femoral, ventral surface.
  12. Shape of the closing surface of the forefemur : (0) straight; (1) proximally curved.
  13. Shape of the tibia : (0) smoothly curved; (1) straight (does not touch the entire surface of the femur when closed); (2) curved (making contact with the femur along its entire surface).
  14. Tibial spur : (0) present; (1) absent.
  15. Prostrate setae on tibial, ventral keel : (0) present; (1) absent.
  16. Foretarsus condition on males and females : (0) 5-segmented in both sexes; (1) 4-segmented in both sexes; (2) sexually dimorphic, 4-segmented on males and 5-segmented on females.
  17. Forebasitarsus morphology : (0) cylindrical; (1) with distal, ventral process; (2) with lanceolate process.
  18. Length of lanceolate process relative to the eutarsus (when present) : (0) short, reaching the apex of the third tarsomere; (1) long, reaching the middle of the fourth tarsomere.
  19. Number of rows of specialized setae on ventral surface of forebasitarsus : (0) two; (1) one; (2) absent.
  20. Foretarsal Stitz organ : (0) absent; (1) present.

Mid- and hind leg

  1. Shape of the hind tibia : (0) unmodified (thin and cylindrical); (1) remarkably expanded (oar-shaped) or fusiform.

Wings

  1. Wings shape : (0) elongate and narrow; (1) broadly oval.
  2. Vesicae : (0) absent; (1) present.

Forewing

  1. Pterostigma shape : (0) fusiform; (1) oval; (2) rectangular; (3) trapezoidal.
  2. Pterostigma color pattern : (0) variegated; (1) transversally divided into two colors; (2) dark with pale medial area.
  3. Costal space width at basal region : (0) slightly expanded; (1) narrow (i.e., C and Sc veins nearly subparallel); (2) notably expanded.
  4. Number of trichosors between the apex of two longitudinal veins on posterior wing margin : (0) 0; (1) 1; (2) ≥ 2.
  5. Forked Subcostal veinlets : (0) absent; (1) present.
  6. Number of crossveins in the subcostal space before pterostigma : (0) 2; (1) 3; (2) 1.
  7. Humeral vein : (0) simple or forked; (1) branched.
  8. Subcostal vein : (0) not fused to RA at pterostigma level; (1) fused to RA at pterostigma level.
  9. Number of crossveins in the radial space : (0) 3; (1) 4; (2) 1; (3) 2.
  10. Shape of the sub-stigmal cell of the radial space : (0) gently curved; (1) oblique; (2) straight; (3) sigmoid; (4) arched.
  11. Shape of the rm1 cell : (0) bar-shaped; (1) arrow-shaped; (2) trapezoidal.
  12. Size of the rm1 cell (if trapezoidal) in relation to M divergence from R : (0) small (i.e., the M vein diverges from R close to RP stem origin); (1) enlarged (i.e., the M vein diverges from R far from RP stem origin).
  13. M and R veins : (0) running parallel and very close at wing base; (1) fused at wing base.
  14. M fork : (0) well beyond to R fork; (1) opposite or nearly opposite to R fork.
  15. Shape of the Cubitus anterior (CuA) : (0) gently curved or only slightly bent near the middle; (1) forming a well-defined obtuse angle.
  16. Cubitus posterior (CuP) of forewing : (0) not approaching A1; (1) approaching or touching A1.
  17. Cubitus posterior (CuP), fork position : (0) slightly before 1m-cu level; (1) at midpoint between 1m-cu and RP origin; (2) opposite to RP origin level.
  18. First anal vein (A1) : (0) basally forked; (1) branched; (2) simple; (3) distally forked.
  19. Second anal vein (A2) : (0) branched; (1) forked.
  20. Third anal vein (A3) : (0) close to jugal lobe; (1) forming a loop with the A2.

Hind wing

  1. Hind wing : (0) nearly as long as the forewing; (1) reaching the distal margin of the forewing pterostigma; (2) reaching the proximal margin of the forewing pterostigma.
  2. Costal and subcostal veins : (0) running subparallel, not fused; (1) fused at or slightly beyond the level of R fork.
  3. RP branches : (0) terminating along posterodistal margin of wing; (1) terminating at wing apex.
  4. Shape of 1r-m (i.e., first crossvein of the radiomedial space) : (0) straight; (1) sigmoidal.
  5. Crossvein connecting the 1r-m to M stem distally : (0) absent; (1) present.
  6. Shape of the media vein (M) : (0) forming angles at its forks and at connection with crossveins; (1) tuning fork-shaped.
  7. First branch of Cubitus anterior (CuA) : (0) simple; (1) forked; (2) candelabrum-shaped.
  8. Cubitus posterior (CuP) : (0) distally forked (i.e., close to posterior wing margin); (1) deeply forked.
  9. Cubitus posterior (CuP) forming a loop with CuA : (0) absent; (1) present.
  10. Intracubital crossvein : (0) absent; (1) subparallel to longitudinal wing axis; (2) reclined, nearly transverse.
  11. Number of crossveins in the cubitoanal space : (0) 0; (1) 1; (2) 2.
  12. Anterior branch of the first anal vein (A1) on hind wing : (0) ending on posterior wing margin fused to CuP; (1) fused to CuP for a short distance; (2) not fused to CuP.

Abdomen

  1. Ornamentation on abdominal tergites : (0) absent; (1) with keeled processes on tergites, including tergites V–VI or V–VII; (2) with keeled processes on tergites, but not including V, VI, and VII.

Male genitalia

  1. Sternites VIII and IX : (0) not fused; (1) fused.
  2. Sternite IX, shape : (0) narrow, with rounded posterior margin; (1) approximately rhomboid, forming a broad canal; (2) pentagonal to semi-triangular with straight vertexes; (3) pentagonal with rounded posterolateral corners; (4) trapezoidal; (5) broadly rounded.
  3. Posteromedial, dorsal canal on sternite IX : (0) absent; (1) present.
  4. Gonocoxites IX : (0) equipped with hook-shaped processes; (1) fused to gonocoxites XI; (2) equipped with digitiform processes; (3) filiform, lacking digitiform processes; (4) greatly reduced and fusiform; (5) bar-shaped with blade-shaped or pointed apex.
  5. Ventral region of the ectoproct : (0) with ventromedial lobe; (1) unmodified; (2) equipped with a patch of stout setae; (3) equipped with anterior lobe and ventromedial curved groove; (4) equipped with anterior bulging area and posterior concave surface; (5) equipped with anterior, flattened lobe anteriorly covered with conical, translucent setae.
  6. Modified setae on posterior surface of the ectoproct : (0) absent; (1) present.
  7. Gonapophyses X : (0) indistinct; (1) paired, not fused to gonocoxites X.
  8. Gonapophyses X (when not fused with gonocoxites X) fused at posterior apex, forming a process : (0) absent; (1) present.
  9. Gonocoxites X (when not fused with gonapophyses X) in lateral view : (0) only slightly wider on anterior apex; (1) narrow, subparallel sided; (2) with anterior apex expanded and dorsally bent; (3) with anterior region conspicuously expanded and dorsally curved or bent.
  10. Gonostyli X : (0) short; (1) long (i.e., whip-shaped).
  11. Gonostyli X, overall shape : (0) dorsally coiled; (1) medially recurved posteriorly; (2) forming an apical incurvation; (3) anteriorly forming a single loop approximately at midlength; (4) anteriorly coiled, forming two loops; (5) dorsally directed and ventrally recurved.
  12. Median lobe of gonocoxites XI : (0) as a transverse, straight bar; (1) as a projection with median notch; (2) as a flattened rounded or quadrangular lobe; (3) with anterior and posterior, lateral, flattened processes; (4) complex and enlarged, with dorsal and ventral part.
  13. Ventromedial posterior area of the gonocoxites XI median lobe (if the median lobe is enlarged and complex) : (0) not developed; (1) convex or protuberant; (2) with prominent posteroventrally projected process; (3) with digitiform or enlarged, caudally projected process.
  14. Ventral ornamentation of the gonocoxites XI median lobe (if the median lobe is enlarged and complex) : (0) absent; (1) a concave covering that forms a curved process; (2) a prominent hook-shaped process; (3) a curved process with bilobed apex; (4) a short and blunt process; (5) an enlarged, curved, and rugose process.

Female genitalia

  1. Tergite IX and ectoproct : (0) not fused; (1) fused.
  2. Gonocoxites VII : (0) unpaired, forming a sternite-like sclerite; (1) paired, as two lateral plates; (2) as lateral plates connected through narrow posterior or median bridge.
  3. Crumena : (0) present; (1) absent.
  4. Gonocoxites + gonapophyses VIII : (0) paired; (1) unpaired, forming a simple sclerite; (2) unpaired, forming a complex structure.
  5. Gonapophyses VIII medial part shape (when forms a complex structure) : (0) forming a tubular projection; (1) forming a broad, box-shaped structure; (2) forming a broad or straight canal; (3) keel-shaped; (4) as a polygonal, convex plate; (5) chamber-shaped.
  6. Bilobed posteromedian process of the gonapophyses VIII (when forming a complex structure) : (0) absent; (1) thickened, with short lobes; (2) elongated and curved; (3) tiny, with short lateral lobes; (4) elongated, Y-shaped.
  7. Tergite IX : (0) continuous with gonocoxites IX; (1) articulated with gonocoxites IX.
  8. Gonocoxites IX, overall shape : (0) ovoid; (1) with hypocauda; (2) hose-shaped.
  9. Gonapophyses IX : (0) absent; (1) present as two tiny sclerites concealed on the inner surface of the gonocoxites IX base.
  10. Callus cercus : (0) present, protuberant; (1) absent.
  11. Bursa copulatrix, sclerotization : (0) completely strongly sclerotized; (1) entirely membranous to slightly sclerotized; (2) with sclerotized and membranous areas.
  12. Distal section of the spermatheca : (0) unmodified, as wide as proximal and medial sections; (1) thin, with blunt diverticulum; (2) conspicuously expanded, sac-shaped; (3) slightly wider than proximal and medial sections, lacking diverticulum; (4) wider than proximal and medial sections, equipped with blunt diverticulum; (5) with conical or trumpet-shaped invagination.
  13. Fertilization canal duct : (0) short; (1) long.
  14. Overall shape of the fertilization canal duct : (0) sinusoidal; (1) curved; (2) spiral-shaped; (3) sinusoid with proximal, triangular, concave portion.

Results

Phylogeny of Symphrasinae

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. 3). The unequivocal changes supporting this clade are the following: head characters such as the diamond-shaped head in frontal view (char 2:3), the pentagonal frontal sclerite (char 3:1), and the presence of hypostomal bridge (char 9:1). Thoracic features include the procoxal insertion approximately at mid-length of prothorax (char 12:2), and the presence of three uneven raised areas on the pronotum (char 16:2). Synapomorphies on the foreleg include the robust forefemur with convex anterior and posterior surfaces (char 21:2), the carinate distal ½ of the posteroventral row of processes of femur (char 22:2), the anteroventral row of processes reduced to proximal region and apex (char 23:2) with the proximal-most process a primary and curved process (char 24:2), presence of thickened setae with globular base adjacent to rows of processes of forefemur (char 28:1), the proximally curved closing surface of forefemur (char 30:1), and the curved tibia (char 31:2). Venational synapomorphies include the presence of more than two trichosors on the posterior wing margin between the apex of two longitudinal veins (char 45:2), the Sc vein fused to RA at pterostigma level (char 49:1), the presence of two crossveins in the radial space of FW (char 50:3), the C and Sc veins fused at or slightly beyond the level of R fork on HW (char 63:1), and the intracubital crossvein of the HW subparallel to longitudinal wing axis (char 71:1). Moreover, genital features supporting the monophyly of Symphrasinae in males are: gonocoxites IX set with apical, digitiform processes (char 78:2), the ventral region of ectoproct set with a patch of stout setae (char 79:2), the paired gonapophyses X not fused to gonocoxites X (char 81:1), the gonostyli X forming an apical incurvation (char 85:2), and the flattened, rounded or quadrangular median lobe of gonocoxites XI (char 86:2). Meanwhile, in the female genitalia, the absence of callus cercus (char 98:1) was recovered as a synapomorphy of the subfamily.

Figure 3. 

Morphological phylogeny of Symphrasinae. Strict consensus of 33 mot parsimonious trees resulting from a New Technology Searching and equal weight.

The genus Anchieta was recovered as monophyletic with A. fasciatellus (Westwood, 1867) sister to the rest of the species (Fig. 3). Synapomorphies for this clade include the rectangular frontal sclerite (char 3:2), a flattened pronotum with short, raised posterior margin (char 16:3), the presence of a blunt process on anterior surface of foretrochanter (char 19:1), the presence of single, sub-basal, primary process on the posteroventral row of processes of forefemur (char 25:2), the rectangular pterostigma of forewing (char 42:2), a single crossvein in the cubitoanal space of HW (char 72:1), the conspicuously expanded distal section of the spermatheca (char 100:2), and the spiral-shaped fertilization canal duct (char 102:2). The fused male sternites VIII and IX (char 75:1) also support the monophyly of Anchieta, yet this character is homoplastic in Mucroberotha vesicaria Tjeder, 1968. Anchieta fumosellus (Westwood, 1867) was recovered as sister of the remaining species of the genus based on the straight substigmal cell of the FW (char 51:2), the hind wing reaching the level of the proximal margin of pterostigma of the forewing (char 62:2), the reclined intracubital cell of HW (char 71:2), and the gonocoxites X with anterior region conspicuously expanded and dorsally curved or bent (char 83:3). The bee-mimicking species of Anchieta form a clade supported by the presence of an expanded or fusiform hind tibia (char 39:1) and the presence of keeled processes on abdominal tergites, but not including those of segments V, VI, and VII (char 74:2) as unequivocal changes (Fig. 3). Within this group of species, a sister group relationship between A. remipes (Gerstaecker, 1888) and A. tinctus Ardila-Camacho & Contreras-Ramos, sp. nov., is supported by the greatly reduced and fusiform male gonocoxites IX (char 78:4), the gonapophyses X fused at posterior apex, forming a process (char 82:1), and the dorsally directed and ventrally recurved gonostyli X (char 85:5). Moreover, A. nebulosus Ardila-Camacho & Machado, sp. nov. and A. nothus (Erichson, 1830) were recovered as sister species based on the presence of keeled processes on abdominal tergites, including tergites V–VI or V–VII, and the gonapophyses medial part of female forming a box-shaped structure (char 93:1).

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. 3). Genitalic features supporting this group include the presence of posteromedial, dorsal canal on sternite IX (char 77:1), the ventral region of ectoproct set with anterior lobe and ventromedial groove (char 79:3), gonostyli X anteriorly coiled, forming two loops (char 85:4), and median lobe of gonocoxites XI complex and enlarged, with dorsal and ventral part (char 86:4).

The genus Trichoscelia was rendered monophyletic with strong support, although the relationships within the genus were largely unresolved and recovered in big polytomies (Fig. 3). As these results are difficult to explain, these will not be used for discussion. The characters supporting the monophyly of this genus include the broadly ovoid palpimacula (char 8:2), the posteroventral row of thickened setae with globular base present on nearly the entire femoral closing surface (char 29:0), the Cubitus anterior (CuA) of FW forming a well-defined obtuse angle (char 56:1), the presence of a crossvein connecting the 1r-m to the M stem distally on the HW (char 66:1), and the ventral region of male ectoproct equipped with anterior bulging area and posterior concave surface (char 79:4). Monophyly of this genus is also supported by the following homoplastic characters: the vertexal region above compound eyes approximately with the middle of the distance between upper limit of compound eyes and frontoclypeal ridge (char 1:2), the enlarged compound eyes (~0.75 of interocular distance at toruli level) (char 4:0), antennal scape > 2 × as long as wide (char 6:2), basal flagellomeres 2–3× as long as wide (char 7:2), prothorax 1.5 × as long as wide (char 13:2), the posteroventral row of processes of forefemur nor raised (char 22:0) and the anteroventral row fully developed (char 23:1) and with proximal-most process not differentiated from other processes (char 24:3). Furthermore, the short lanceolate process of forebasitarsus (i.e., reaching the apex of third tarsomere) (char 36:0) and the bursa copulatrix with sclerotized and membranous areas (char 99:2) (homoplastic in P. insolita Ardila-Camacho & Contreras-Ramos, sp. nov. and Plega, respectively) also support the monophyly of Trichoscelia. The conical or trumpet-shaped invagination on distal section of the spermatheca was recovered as a synapomorphy of all Trichoscelia species exclusive of T. wintertoni Ardila-Camacho & Contreras-Ramos, sp. nov., nonetheless it was an ambiguous character state in this latter species, so this feature should be considered as a synapomorphy of the whole genus.

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. 3). In this clade, P. zikani Navás, 1936 was recovered as the sister species of the rest of the species of the genus which form a clade supported by a synapomorphy, i.e., supra-antennal region set with prominent lateral, setose protuberances (char 10:2), and a homoplasious character state, i.e., a sigmoid sub-stigmal cell of the FW (char 51:3).

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).

Taxonomy

Class Insecta Linnaeus, 1758

Order Neuroptera Linnaeus, 1758

Family Rhachiberothidae Tjeder, 1959

Symphrasinae , Navás, 1909

Symphrasini Navás 1909: 484.

Anisopterinae Enderlein 1910: 341.

Platymantispinae Rehn 1939: 82.

Further description

Penny (1982a); Penny and da Costa (1983); Lambkin (1986a); Wedmann and Makarkin (2007); Lu et al. (2020); Ardila-Camacho et al. (2021a); Li et al. (2023).

Taxonomy

Rehn (1939b); Penny (1982a, b); Penny and da Costa (1983); Ardila-Camacho and García (2015); Ardila-Camacho et al. (2018, 2019).

Key to genera

Navás (1912); Banks (1913); Penny (1982a, b); Penny and da Costa (1983); Hoffman (2002).

Phylogeny

Penny and da Costa (1983) (Symphrasinae sister group of the remaining Mantispidae); Lambkin (1986a) (Symphrasinae sister group of other subfamilies of Mantispidae); Willmann (1990) (Symphrasinae sister group of other subfamilies of Mantispidae); Liu et al. (2015) (Symphrasinae sister group of other subfamilies of Mantispidae); Winterton et al. (2018) (Symphrasinae + Rhachiberothinae sister of Berothidae + (Mantispinae + (Calomantispinae + Drepanicinae)); Shi et al. (2019) (Symphrasinae sister of Mesomantispinae); Shi et al. (2020) (Symphrasinae sister of Doratomantispa Poinar); Lu et al. (2020) (Symphrasinae sister of Doratomantispinae + (Drepanicinae + (Calomantispinae + Mantispinae)); Ardila-Camacho et al. (2021a) (Symphrasinae + Paraberothinae sister of Rhachiberothinae).

Diagnosis

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.

Biology

Summarized in Ardila-Camacho et al. (2021b).

Key to Symphrasinae genera

1 Foretrochanter with blunt process on anterior surface (Fig. 1B); HW with first branch of CuA simple or forked (Fig. 4B); male sternites VIII and IX fused (Fig. 5B); distal portion of spermatheca noticeably expanded, sac-like (Fig. 15H) Anchieta Navás 1909
Foretrochanter without blunt process on anterior surface; HW with first branch of CuA candelabrum-shaped (Fig. 2B); male sternites VIII and IX as separate units; distal portion of spermatheca narrow, generally with a diverticulum or invagination (Figs 27I, 82I) 2
2 Third labial palpomere expanded, with broadly ovoid palpimacula (Fig. 83C); forefemur with rows of thickened setae with globular base extending nearly over the entire closing surface; posteroventral processes row uniform, without spine-shaped processes (Fig. 89E, F); HW with 1r-m distally connected to M stem through a short crossvein (Fig. 81B); distal section of spermatheca with conical or trumpet-shaped invagination (Fig. 84I) 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. 1D); posteroventral processes row with two primary, spine-shaped processes on proximal 1/2 (Fig. 1E); HW with 1r-m not distally connected to M stem through a short crossvein; distal section of spermatheca with or without a diverticulum (Figs 43I, 78B) Plega Navás, 1928

Anchieta Navás, 1909

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 Penny (1982b): 216.

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 Rehn (1939a).

Anchieta Bechyné, 1954: 176 (in Coleoptera). Junior homonym of Anchieta Navás, 1909.

Further description

Gerstaecker (1888): 117; Enderlein (1910): 375 (as Anisoptera); Navás (1912): 201; Penny (1982b): 417; Penny and da Costa (1983): 610.

Taxonomy

Gerstaecker (1888): 117 (Anisoptera = Trichoscelia); Enderlein (1910): 376 (Anchieta = Trichoscelia); Navás (1912): 201 (Anchieta valid genus); Banks (1913): 205, 206 (Anchieta and Trichoscelia = Anisoptera).

Key to species

Banks (1913): 207–208 (Westwood’s species); Penny (1982a): 418; Penny and da Costa (1983): 611.

List of species

Penny (1977): 37 (as Trichoscelia).

Diagnosis

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.

Description

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.

Included species

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)

Biology

Summarized in Ardila-Camacho et al. (2021b).

Etymology

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 Navás (1909).

Key to the species of Anchieta

1 Hind tibia narrow and unmodified or only slightly widened (Fig. 10A) 2
Hind tibia noticeably expanded and flattened, fusiform or oar-shaped (Fig. 4F) 4
2 Body bright orange with dark brown bands (Fig. 8A); male gonocoxite IX with 8–13 apical processes arranged as a brush (Fig. 9C) 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. 11C–E); female gonapophyses VIII forming a tubular projection (Fig. 11H) Anchieta fumosellus (Westwood, 1867)
Male gonocoxite IX filiform, with posterior apex sharply pointed (Fig. 23C–E); female gonapophyses VIII forming a chamber-shaped structure (Fig. 23H) Anchieta sophiae Ardila-Camacho & Contreras-Ramos, sp. nov.
4 Body nearly completely dark brown (Fig. 14A) 5
Body mostly pale orange, with brown marks (Fig. 4A) 7
5 Male gonocoxite IX reduced, fusiform and helical, ventrally attached to lateral arms of gonocoxites XI (Fig. 25C–E); female gonapophyses VIII medial part composed by quadrangular plates forming a canal (Fig. 25G) Anchieta tinctus Ardila-Camacho & Contreras-Ramos, sp. nov.
Male gonocoxite IX well-developed, filiform or thickened with digitiform processes (Figs 13D, 15C); female gonapophyses VIII medial part box-shaped (Fig. 13H) 6
6 Male gonocoxite IX with posterior apex ventrally curved, and set with three short processes (Fig. 15C–E) Anchieta nothus (Erichson, 1830)
Male gonocoxite IX filiform, lacking processes (Fig. 13 C–E) Anchieta nebulosus Ardila-Camacho & Machado, sp. nov.
7 Forewing costal space narrow (Fig. 6B); hind tibia fusiform (Fig. 6F); male gonocoxite IX with posterior apex pointed and set with four short, preapical, spine-shaped processes, located on the ventral surface (Fig. 7C–E) Anchieta bellus (Westwood, 1867)
Forewing costal space medially expanded (Fig. 4B); hind tibia noticeably expanded, oar-shaped (Fig. 4F); processes on male gonocoxite IX with different arrangement 8
8 Male sternite IX with broad posteromedial, quadrangular lobe (Fig. 19B); male gonocoxite IX reduced and fusiform (Fig. 19A); male gonostyli X dorsally recurved and anteroventrally projected (Fig. 19A) Anchieta remipes (Gerstaecker, 1888)
Male sternite IX approximately rhomboid, with broad, rounded, posteromedial canal (Fig. 17E); male gonocoxite IX well-developed, bearing digitiform processes (Fig. 17C); male gonostyli X anteroventrally recurved, forming and apical incurvation in lateral view (Fig. 17C) 9
9 Male gonocoxite IX with posterior apex strongly lateroventrally recurved, set with two apical and four preapical, short processes (Fig. 17C–E) 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. 21C–E); male gonocoxites X anterior apex not expanded, laterally flattened (Fig. 21C, D) Anchieta romani (Esben-Petersen, 1817)
Male gonocoxite IX apex ventrally curved, set with one short, preapical process (Fig. 5C); male gonocoxites X anterior apex noticeably expanded, dorsally recurved (Fig. 5C, D) Anchieta apiculasaeva Thouvenot, 2009

Achieta apiculasaeva Thouvenot, 2009

Figs 4, 5

Anchieta apiculasaeva Thouvenot, 2009: 226. Holotype: male, French Guiana, [Régina] Piste de Kaw (MHNL), high resolution images examined.

Material examined

Holotype. French Guiana • ♂; [Régina] Route de KAW pk 3; 03 Mar. 1981; G. Tavakilian leg.; MHNL.

Other material

Brazil • 1 ♂; abdomen cleared and stored in a microvial; INPA. – Mato Grosso • 1 ♂; Novo Mundo, Pq. Est. do Cristalino; 9°27'6.12"S, 55°50'22.56"W; 21–25 Jun. 2007; R.R. Cavichioli and A.C. Domahovski leg.; light trap; DZUP. French Guiana • 1 ♀; Matoury; Mont Matoury, Mont Grand Matoury; 4°51'31.00"N, 52°22'36.00"W; 28 Sep. 2014; flight interception trap, Colline entourée de savanes et zones dégradées; CSCA.

Diagnosis

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.

Description

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. 4). Head. Mostly orange, vertexal region with broad, dark brown, transverse band, laterally with rows of dark brown setae; supraantennal area with lateral brown triangular markings located above the toruli, such areas connected to the frontal markings through the interantennal area; frons with brown lateral semi-triangular markings located below the toruli, forming an inverted V-shaped pattern. Antennal scape orange ventrally, brown dorsally, pedicel brown, flagellum brown, except for four or five yellow preapical flagellomeres. Clypeus yellow with brown suffusions, labrum with brown central region; mandible dark amber; maxillary palpus pale brown; labium orange; labial palpus with first and fourth palpomeres orange, second, third and fifth brown, palpimacula pale brown. Thorax. Pronotum dark brown with an orange band on distal 1/3, anterior margin dark brown, with concolorous setae. Meso- and metanotum mostly orange, with brown markings on area adjacent to parapsidal sutures, and on scutellum, setae pale brown; episternum dark brown; postfurcasternum dark brown with small, posterior, yellow area. Pteropleura mainly orange with some pale brown areas, setation orange. Foreleg. Coxa yellow at base, apex, and anterior surface; posterior surface with an extensive dark brown mark; setae mostly concolorous, except on anterior surface with interspersed orange and dark brown setae; trochanter bicolor, dorsally pale brown, ventrally orange, with interspersed brown and orange setae. Femur mainly orange with extensive, dark brown, dorsal mark, extending to anterior surface; posterior surface with two elongated brown markings, one located at apex, the other at middle; with interspersed orange and brown setae; tibia bicolor, basal ½ orange, with brown mark on dorsal surface; distal ½ brown, clavate setae orange; basitarsus pale brown with pale amber apex, remaining tarsomeres pale brown. Mid and hind legs. Mid-leg with predominantly yellow coxa, trochanter brown, femur orange; tibia predominantly pale brown; tarsus orange with dark brown setae on ventral surface. Hind leg coxa orange with pale brown suffusions, trochanter pale brown, femur orange with pale brown, medial suffusion; tibia orange with pale brown posterior margin; tarsus orange with dark brown setae on ventral surface. Wings. Forewing hyaline with orange areas base and area adjacent to proximal region of pterostigma; an orange transverse band located at R fork; pterostigma bicolor, pale brown on proximal ½, orange on distal ½; venation predominantly orange with brown suffusions at wing base and base and middle of R +M; apex of RP and M branches pale brown. Hind wing hyaline, base of subcostal space orange; pterostigma orange with blackish setation; venation orange with blackish suffusions at wing base. Abdomen. Tergites yellow with brown, irregular areas, tergite IV completely yellow; sternites yellow with orange setae; pleural membrane yellow on proximal 1/3 of abdomen, brown in the rest.

Figure 4. 

Anchieta apiculasaeva Thouvenot, 2009 A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, posterior surface F hind tibia, outer surface.

Morphology (Fig. 4). Head. Diamond-shaped in frontal view, vertexal region domed above compound eyes, rugose, laterally with rows of short, reclined setae; paraocular area concave; coronal suture distinct. Compound eye hemispherical, as wide as ½ the interocular distance at toruli level. Antenna moniliform, flagellum slightly dorsoventrally flattened, with 50 flagellomeres, discoidal in shape, except for those of apex, narrower; all flagellomeres covered with medial ring of short setae. Thorax. Pronotum nearly as long as wide, with groove contiguous to lateral and distal margins; posterior margin with slight outgrowth in lateral view; entire surface with abundant, thick setae arising flush the pronotal surface; episternum with short, thin setae; postfurcasternum quadrangular. Mesonotum slightly wider than long, with abundant long, thickened setae on the medial region; metanotum ~ 3× as wide as long, glabrous. Pteropleura covered with abundant long, thin setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, with abundant fine and long setae; trochanter semi-triangular, densely setose, dorsally with long, thickened setae, anterior surface with blunt process. Robust femur, with abundant long and fine setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes, with conical Stitz organs (process/seta ratio 1:1 or 2:1), proximally with a more developed, sub-basal, spine-shaped process (process/seta ratio 4:1); adjacent row of thickened setae with globular base present on distal ½; anteroventral row of integumentary specializations reduced to proximal region and apex, composed of tubercle-shaped processes with conical setae; primary process present, spine-shaped, curved; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, with a row of prostrate setae on the closing surface, and patch of clavate setae apically on anterior surface. Basitarsus with long lanceolate process, proximal region with clavate setae on anterior surface, and a row of prostrate setae ventrally. Mid and hind legs. Mid-leg covered with abundant fine and long setae, tibia slightly widened. Hind leg densely covered with long, fine setae, except on the tarsus, where they are shorter, tibia notably widened and laterally flattened, oar-shaped. Wings. Forewing oval, venation densely setose, trichosors present along wing margin; costal space slightly widened medially, humeral vein simple, 10–12 veinlets present; pterostigma rectangular; subcostal space with a single medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge with the RA; rarp2 straight with three veins arising from it, three or four veins from rarp1; M fused basally to R; RP base located near separation of M and RA, M fork near such separation; 1r-m located between RP base and M fork, forming a small trapezoidal cell; five or six gradate crossveins present; Cu deeply forked, CuP basally angled, forked; A1 simple, A2 distally forked. Hind wing oval, notably smaller and narrower than forewing; costal space narrow and reduced, with seven veinlets; C and Sc fused at proximal 1/3 of wing length, subcostal space without crossveins; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge with RA; pterostigma elongated, slightly widened distally; radial area widened with single, sinuous crossvein; two veins arising from rarp1, two from rarp2. M vein forked at level of R fork; Cu deeply forked, CuA sinuous, distally forked; CuP distally, anteriorly bent, with three branches; A1 arched, A2 short, simple. Abdomen. Medially widened, tergites III and IV with short, keeled processes, tergites V–IX with abundant long, thin setae; sternites densely setose.

Male genitalia (Fig. 5A–E). Tergite IX medially narrower than laterally, lateral region covered with abundant long and thickened setae. Sternites VIII and IX fused, with weakly marked fusion line; sternite VIII subrectangular with abundant long and thickened setae; sternite IX U-shaped, with slightly concave lateral margins, and abundant long and thickened setae anteriorly and laterally; posteromedially with convex and glabrous area. Gonocoxite IX short, sinuous, posterior apex ventromedially gently curved, with two processes, one apical, and a shorter preapical, sometimes a third, more proximal, and shorter is present. Ectoproct ovoid, setose, posteroventrally with an area covered with 17 or 18 stout setae; outer surface with smooth, glabrous, bulging area; inner surface with sclerotized ventral area, set with a patch of microtrichia. Gonocoxites X thickened, medially slightly constricted, hourglass-shaped, ventrally canaliculated; anterior ½ expanded, gently curved dorsally, posterior ½ triangular, with paired lateral and dorsal processes. Gonostyli X base triangular, thickened, with curved lateral processes, dorsally concave, the rest of the structure whip-shaped, short, distal portion curved. Gonapophyses X rod-shaped, straight, narrow, anterior apex spatulate, posterior apex curved dorsally; gonapophyses forming a V-shaped structure joined by membranes. Gonocoxites XI thin, U-shaped, medial lobe flattened, weakly sclerotized at center, posterior margin almost straight, lateral borders blunt; lateral arms of gonocoxites straight, apex ventromedially curved.

Figure 5. 

Anchieta apiculasaeva Thouvenot, 2009 A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G gonapophyses VIII of female, ventral.

Female genitalia (Fig. 5F, G). Sternum VII (gonocoxites VII) composed of two medially fused trapezoidal lateral plates, posterior margin obtuse. Tergite VIII narrower medially than laterally, enclosing the segment spiracle, lateral margin triangular. Gonocoxites VIII as two concave, medially fused, trapezoidal plates; gonapophyses VIII medial part ventrally projected forming a broad canal, distal margin rounded, ventrally curved; lateral part of gonapophyses VIII a trapezoidal plate, forming a chamber-shaped plate, dorsally with two convex areas, located behind the genital pore. Tergite IX + ectoproct semi-triangular. Gonocoxite IX long, straight, narrow. Bursa copulatrix unsclerotized, long, conical; spermatheca spiral-shaped; proximal section long, thin, progressively thickened, spiral-shaped, forming three coils; medial section slightly narrower than thickest part of proximal section, entangled, forming several convolutions; distal section thicker than proximal and medial sections, progressively wider towards the apex. Fertilization canal duct long, spiral-shaped; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Mato Grosso), French Guiana (Matoury, Régina).

Remarks

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 (Thouvenot 2009). After looking for loan of the holotype in the context of the present revision, M. Thouvenot informed that the type was deposited in the Musée des Confluences at Lyon, France (MHNL) (M. Thouvenot, C. Audibert, pers. comm., 18 June 2019). Unfortunately, because the holotype is the only specimen known of this species, a loan from the Musée des Confluences was not possible, so only high-resolution images of the type were studied. Herein several additional specimens were identified, and the distribution range of this species is extended to Matoury in French Guiana, and to Matto Grosso in Brazil.

This species exhibits the same coloration pattern of other species of the genus mimicking stingless bees of the genus Ptilotrigona Moure, 1951 (Rasmussen and Ardila-Camacho 2021). Based on the wing shape and venation, hind tibia shape, and morphology of the male genitalia, this species appears to be closely related to A. romani. In the phylogeny of the subfamily, this species was recovered within the clade composing all the species exhibiting Batesian mimicry with stingless bees (i.e., A. bellus, A. apiculasaeva, A. romani, A. remipes, A. tinctus, A. partheniellus, A. nebulosus, and A. nothus).

Anchieta bellus (Westwood, 1867)

Figs 6, 7

Mantispa (Trichoscelia) bella Westwood, 1867: 502. Holotype: female, Amazonia (OUMNH), high resolution photos studied.

Mantispa (Trichoscelia) eurydella Westwood, 1867: 501. Holotype: female, Amazonia (OUMNH), high resolution photos studied. New synonym.

Anisoptera bella (Westwood, 1867). Gerstaecker (1888).

Anisoptera eurydella (Westwood, 1867). Gerstaecker (1888).

Trichoscelia bella (Westwood, 1867). Enderlein (1910).

Trichoscelia eurydella (Westwood, 1867). Enderlein (1910).

Anchieta eurydella (Westwood, 1867). Penny (1982a), Ohl (2004), Ardila-Camacho and García (2015), Ardila-Camacho et al. (2018).

Material examined

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; OUMNH.

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; OUMNH.

Other material

Brazil – Amazonas • 1 ♂; Mamiraua, varzea; 03°02'54"S, 64°51'02"W; 22 Sep. 1993; I.S. Gorayeb and O.T. Silveira leg.; MPEG. – Pará • 1 ♀; Tucuruí, Rio Tocantins, Saude; 1–3 Jun. 1984; arm. Suspensa; MPEG; • 1 ♀; Belém, Floresta APEG; 13–16 Aug. 1983; I.S. Gorayeb leg.; MPEG. • 1 ♀; same data as for preceding; 10–14 Jan. 1983; I.S. Gorayeb e equipe leg.; arm. Suspensa; MPEG. • 1 ♀; Serra Norte, estrada do Manganes; 12 May. 1984; arm. Suspensa; MPEG. • 1 ♀; same data as for preceding; N3; 26–29 Jun. 1985; arm. Suspensa; MPEG.

Colombia • 1 ♀; 1997; Anchieta eurydella det. M. Ohl, 2006; ZMB.

Suriname • 1 ♂; Brokopondo; 24 Apr. 1965; G.F. Mees leg.; A. remipes, det. L.A. Stange; FSCA.

Diagnosis

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.

Description

Measurements. Male (n = 1). Forewing length: 8.0 mm; Hind wing length: 4.9 mm.

Coloration (Fig. 6). Head. Mostly orange, vertexal region with transverse brown band connected anteriorly to triangular dark brown mark on supraantennal region through coronal suture, marking of supraantennal area enclosing central orange spot; laterally with orange setae. Antennal scape mainly orange, amber at apex, pedicel pale brown; flagellum brown, except for five orange, preapical flagellomeres. Frons with an inverted, brown, V-shaped marking. Clypeus and labrum with yellowish setae; mandible orange, with amber apex; maxillary palpus orange; labium orange, including palpimacula. Thorax. Pronotum pale brown with transverse orange band on distal 1/3, setae pale brown; episternum; postfurcasternum orange with brown suffusions. Mesonotum orange with pale brown suffusions, medial region of scutum and scutellum brown, setae orange; metanotum orange with dark brown suffusions mainly at center; pre-episternum brown; mesopleuron with episternum mostly orange, epimeron brown, setation orange; metapleuron with orange episternum, epimeron brown, the entire surface with orange setae. Foreleg. Coxa mostly orange with a pale brown area externally on basal ½, this area plus the apex with dark brown setae, the rest of the surface with orange setae; trochanter orange on ventral surface, pale brown on dorsal surface, setae mostly orange, with a few dark brown setae on dorsal surface. Femur orange with pale brown areas on dorsal and lateral surfaces, setae mostly orange with some scattered brown setae; tibia with orange basal ½, distal ½ amber, clavate setae orange; basitarsus orange with amber lanceolate process; second to fourth tarsomere orange. Mid- and hind leg. Mid-leg coxa mostly orange, with pale brown margins and orange setae; trochanter pale brown; femur orange, pale brown on center, with concolorous setae; tibia with basal ½ and tip orange, preapical region pale brown with dark brown setae; tarsomeres orange. Hind leg coxa mostly orange with pale brown margins and orange setae, trochanter pale brown, femur with orange base and apex, pale brown on remaining surface; tibia with orange base and apex, medial region with brown suffusions; tarsus pale brown with orange setae. Wings. Forewing hyaline, pale amber at base, with transverse pale amber band at R fork, and area surrounding the 1ra-rp; proximal ¾ of pterostigma pale brown, orange on distal ¼; venation uniformly pale brown; hind wing mainly hyaline, pale brown at base; base of subcostal area pale amber; pterostigma pale brown, venation uniformly pale brown, with paler areas on cubital and anal veins. Abdomen. Orange with pale brown areas.

Figure 6. 

Anchieta bellus (Westwood, 1867) A female habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, posterior surface F hind tibia, outer surface.

Morphology (Fig. 6). Head. Diamond-shaped in frontal view, smooth; vertexal region domed above compound eyes, with lateral rows of short, reclined setae; paraocular area; coronal suture distinct. Antenna moniliform, flagellum with 45 flagellomeres, discoidal in shape, those at the apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispheric, as wide as ½ the interocular distance at toruli level. Thorax. Pronotum slightly longer than wide, with groove contiguous to lateral and distal margins; in lateral view, posterior margin slightly elevated, entire surface with abundant long and thickened setae, arising flush the pronotal surface; episternum setose; postfurcasternum quadrangular. Mesonotum slightly wider than long, with abundant long and thick setae on medial region; metanotum ~ 3× as wide as long with. Pteropleura covered with abundant fine, long setae. Foreleg. Coxa as long as femur, sub-cylindrical, slightly expanded on preapical region, with abundant long and fine setae; trochanter subconical, densely setose, dorsally with a tuft of long and thick setae, anterior surface with blunt process covered with long setae. Femur robust, with abundant fine and long setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes with conical Stitz organs, proximally with a more developed sub-basal, spine-shaped process, adjacent row of thickened setae with globular base present on distal ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes with conical setae; primary, spine-shaped, curved process present; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as the femur, curved, glabrous, closing surface with a row of prostrate setae; with a patch of clavate setae apically on anterior surface. Basitarsus with long lanceolate process, proximal part with clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid and hind legs. Mid-leg with coxa, trochanter and femur set with long and fine setae, tibia slightly expanded medially, with abundant long and thickened setae; tarsus covered with short setae, the first tarsomere as long as the next three together, the second as long as the third and fourth together. Hind leg with the coxa covered with long, fine setae, trochanter with few long, fine setae; femur cylindrical, covered with long, fine setae; tibia moderately expanded, laterally flattened, fusiform, densely setose; tarsomeres with short, thin setae, the first tarsomere as long as the next three together, the second as long as the third and fourth together. Wings. Forewing oval, venation setose, trichosors present along wing margin; costal space narrow, humeral vein forked, 8–10 crossveins present; pterostigma rectangular; subcostal space with single, medially located crossvein, Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge with the RA; rarp2 straight with three veins arising from it, two from rarp1; M basally fused to R; RP located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork; four or five gradate crossveins present; Cu deeply forked, CuP proximally angled, approaching A1, forked; A1 without simple. Hind wing notably smaller and narrower than forewing, oval; narrow and reduced costal space with five crossveins; C and Sc fused at proximal 1/3 of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma, to merge with RA; subcostal space without crossveins; pterostigma elongated, with the subapical portion expanded; radial space with single, sinuous crossvein; two veins arising from rarp1, one from rarp2. M forked at or slightly beyond R fork; 1r-m sigmoid; gradate crossveins absent. Cu vein deeply forked, CuA sinuous, distally forked; CuP distally anteriorly gently curved near wing margin, branched; A1 arched, A2 short, simple. Abdomen. Medially widened, tergites of abdominal segments III and IV with short keeled posteromedial projections.

Male genitalia (Fig. 7A–E). Tergite IX notably narrower medially than laterally; posterolaterally setose and with small cleft. Sternites VIII and IX fused, with fusion line distinct; sternite VIII ~ 2× as wide as long; sternite IX narrow, ventrally projected, transversely curved, broadly canaliculate, receiving the apex of gonapophyses X. Gonocoxites IX narrow, curved, base spatulate; apex pointed with four short, preapical processes, located on ventral surface. Ectoproct ovoid, lateral surface with smooth protrusion, posteroventral region with ~ 11 short and thick setae; ventral region with moderately sclerotized plate, equipped with few thin and short setae. Gonocoxites X medially constricted, hourglass-shaped, ventrally canaliculate; anterior apex expanded, curved dorsally; posterior apex with dorsal and lateral processes; base of gonostyli X thickened, concave, with lateral rounded lobes, the rest of the structure thin, ventrally curved, whip-shaped, short, reaching anterior margin of sternite VII, with terminal incurvation. Gonapophyses X elongated, straight, narrow, with the anterior apex slightly dorsally bent; gonapophyses arranged in a V-shaped structure, joined by membranes. Gonocoxites XI narrow, U-shaped, medial lobe as a thin, quadrangular plate with blunt corners; lateral arms of gonocoxites XI short with anterior apex curved ventrally.

Figure 7. 

Anchieta bellus (Westwood, 1867) A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca.

Female genitalia (Fig. 7F–H). Sternum VII (gonocoxites VII) rectangular with posterior margin concave. Tergite VIII as wide medially as laterally, enclosing the segment, lateral margin rounded. Gonocoxites VIII as a bar-shaped, concave plate, forming and obtuse angle; gonapophyses VIII medial part, ventrally projected, forming a broad, short, blunt canal; lateral part of gonapophyses VIII a narrow, convex plate, forming a chamber-shaped covering dorsally, located behind the genital pore. Tergite IX + ectoproct triangular. Gonocoxite IX long, straight, narrow, as long as the last six abdominal segments together. Bursa copulatrix unsclerotized, short, funnel-shaped; spermatheca spiral-shaped; proximal section long, thin, progressively thickened, spiral-shaped, forming three coils; medial section slightly thicker than proximal section, forming several convolutions; distal section thicker than proximal and medial sections, progressively wider towards the apex, terminal portion expanded. Fertilization canal duct long, spiral-shaped, forming four coils; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Amazonas, Pará), Colombia, French Guiana (Saint-Laurent-du-Maroni) (doubtful record), Suriname (Brokopondo).

Remarks

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 Penny (1982a) and Penny and da Costa (1983) to separate these species are interpreted as intraspecific variation in the intensity of the coloration of the body (darker in A. eurydella and paler in A. bellus), a rather common attribute in Symphrasinae. Anchieta eurydella has also “Amazons” as collecting data, and the type, like A. bellus is a female specimen. Penny (1982a) and Penny and da Costa (1983) provided records of A. bellus from the Brazilian states of Amazonas and Pará, and from Saint-Laurent-du-Maroni, French Guiana, nevertheless, the taxonomic identification of the specimens from Amazonas and Saint-Laurent-du-Maroni is herein considered as doubtful, primarily the first record which was made from a female. The Brazilian specimen could actually correspond to A. partheniellus, whereas the specimen from French Guiana could be A. apiculasaeva. Further examination of these specimens will be necessary to corroborate these records.

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 (Rasmussen and Ardila-Camacho 2021).

Anchieta fasciatellus (Westwood, 1867)

Figs 8, 9

Mantispa (Trichoscelia) fasciatella Westwood 1867: 503. Holotype: female, Santa Martha, Venezuela (OUMNH), high resolution images studied.

Anisoptera fasciatella (Westwood, 1867). Gerstaecker (1888).

Trichoscelia fasciatella Westwood, 1867. Enderlein (1910).

Plega fasciatella (Westwood, 1867). Penny (1982a), Ohl (2004), Ardila-Camacho and García (2015).

Anchieta fasciatella (Westwood, 1867). Ardila-Camacho et al. (2018).

Material examined

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; OUMNH.

Other material

Colombia – Bolívar • 1 ♂; Mompox; 09°14'N, 74°25'W; 33 m; Dec. 1994; MEFLG. • 1 ♂; Zambrano, Hda. Monterrey; 9°45'N, 74°49'W; 70 m; 14 Aug. 1993; F. Fernandez; Malaise No. 6, Lata suelo; ICN. – Cesar • 1 ♀; Chiriguana District, lake Sapatoza región; 08 Sep. 1924; C. Allen, BMNH(E) 1201805, Brit Mus. 1925-576, NHMUK 013802797 Trichoscelia fasciatella Westwood; NHMUK. • 1 ♂; same data collection as for preceding; BMNH(E) 1201814, Brit. Mus. 1925-576, NHMUK 013802798, Trichoscelia fasciatella Westwood; NHMUK. • 1 ♂; Colombia; Mar. 1968; Saunders leg.; BMNH(E) 1201812, NHMUK 013802800; NHMUK. – Magdalena • 1 ♀; Aracataca; 21 Apr.; Darlington leg.; MCZ • 1 ♀; Neguanje, PNN Tayrona; 11°20'N, 74°02'W; 10 m; 21 Feb.–5 Mar. 2001; R. Henriquez leg.; Malaise m–1351; ICN. – Santander • 1 ♀; Puerto Parra, Campo Capote; 06°37'08.6"N, 73°54'30"W; 146 m; C. Sarmiento leg.; light trap; ICN.

Panama – Canal Zone • 1 ♂; Albrook Forest Site; 30.40 m; 25–26 Jul. 1968; R. S. Hutton leg.; black light trap; UAAM • 1 ♂; same data as for preceding; 13–14 Jul. 1967; Hutton & Llaurudo leg.; UAAM • 2 ♂; same data as for preceding; 24–25 Aug. 1967; Hutton & Llaurudo leg.; black light trap; UAAM • 1 ♂; same data as for preceding 29 Feb.–1 Mar. 1968; R.S. Hutton leg.; UAAM • 2 ♂; same data as for preceding; 3–4 Aug. 1967; Hutton & Llaurudo leg.; UAAM. – Darién • 1 ♂; Patino R. Pan; 18 Jul. 1952; F.B. Blanton leg.; det. L.A. Stange; FSCA. – Panamá • 1? 1♂; 7–10 Km. N El Llano; 14–22 May. 1993; E. Giesbert leg.; det. L.A. Stange; FSCA • 1 ♂; same data as for preceding; 16–22 May. 1987; E. Giesbert leg.; FSCA • 1 ♂; same data as for preceding; B-10 Km N. El Llano; 26 Apr.–4 May. 1992; E. Giesbert leg.; FSCA • 1 ♂; same data as for preceding; 8–10 km N El Llano; 927.696, -7.896.416; 26 Apr.–04 May. 1992; E. Giesbert leg.; TAMUIC • 2 ♂, 7.5–13 Km El Llano; 21–22 May. 1994; F.T. Hovare leg; CAS • 1 ♂; Panama; 1924; L.E. Cheesman leg.; BMNH(E) 1201813, St. George Exp. B.M. 1925-573, NHMUK 013802799, Trichoscelia fasciatella Westwood; NHMUK. – Panamá Oeste • 1 ♂; Barro Colorado Island; 18 Apr. 1874; Duckworth leg.; USNM • 2♂; Barro Colorado Island; 3–9 Apr. 1985; H. Wolda leg.; Ex. Blacklight; UCD • 1 ♂; Barro Colorado Island; 17–23 Apr. 1985; H. Wolda leg.; UCD.

Diagnosis

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.

Description

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. 8). Head. Dark reddish brown to black, vertexal region with lateral rows of black setae; postgena dark reddish brown. Antenna black. Mandible with basal ½ black, distal ½ amber; maxillary palpus black; labium dark reddish brown, palpimacula brown. Thorax. Pronotum orange, with interspersed black and orange setae. Episternum and postfurcasternum orange. Meso- and metanotum orange, with interspersed orange, brown and black setae; pteropleura completely orange. Foreleg. Coxa completely orange; trochanter orange or dark reddish brown ventrally and brown dorsally, with concolorous setae. Femur orange; tibia with basal ¼ orange, the rest brown to dark reddish brown, apex generally brown, clavate setae yellowish; basitarsus amber with black tip; second to fourth tarsomere pale brown. Mid- and hind legs. Mid-leg with coxa, trochanter and femur orange; tibia bicolor, basal ½ orange, distal ½ dark reddish brown to black; tarsus dark reddish brown, ventrally with rows of black setae on the distal margin. Hind leg with coxa, trochanter and femur orange; tibia with basal ¼ orange, the rest dark reddish brown to black; tarsus dark reddish brown, ventrally with rows of black setae. Wings. Forewing pale orange, dark amber bands at mid length and apex; pterostigma varying with blackish amber from proximal margin to the middle, the rest orange. Venation mainly orange, brown on middle and apical ¼. Hind wing pale orange, subcostal space apex and wingtip amber; pterostigma bicolor, proximal ½ to ¾ blackish amber, orange on the rest. Venation mainly brown, basal ½ of C, Sc, RA, RP veins, M base and basal ½ of Cu orange; apex of C+Sc and RA beyond pterostigma orange. Abdomen. Basal ½ of the abdomen orange; tergite V bicolor, orange on proximal region, dark brown on region; tergites of abdominal segments VI to ectoproct black; sternites of abdominal segments VI–IX black.

Figure 8. 

Anchieta fasciatellus (Westwood, 1867) A male habitus, dorsal B wings C head, dorsal D pronotum, dorsal E forefemur, anterior surface F forefemur, posterior surface.

Morphology (Fig. 8). Head. Diamond-shaped in frontal view, smooth, vertexal region domed over compound eyes, paraocular area concave; coronal suture discrete. Compound eye hemispheric, as wide as 1/2 of interocular distance at toruli level. Antenna moniliform, flagellum with 58–63 flagellomeres, discoidal in shape, those of the distal ½ wider, except the apical ones, narrow; all flagellomeres with medial ring of short setae. Thorax. Pronotum slightly longer than wide, with shallow medial concavity; in lateral view, posterior region slightly elevated, entire surface with interspersed thin and thick setae that arise flush the pronotal surface. Postfurcasternum semicircular. Mesonotum ~ 2× as wide as long, with a few fine setae; metanotum ~ 3× as wide as long, mostly glabrous. Pteropleura covered with few fine setae. Foreleg. Coxa slightly shorter than femur, cylindrical, slightly expanded on preapical region, with fine and short setae; trochanter subtrapezoidal, setose, anterior surface with conspicuous, blunt process; femur robust, densely covered with short and thin setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of thickened tubercle-shaped processes with conical Stitz organs, proximally with a more developed sub-basal, spine-shaped process; adjacent row of thickened setae with globular base reduced to a single apical seta; anteroventral row of integumentary specializations reduced to proximal region and apex, composed of tubercle-shaped processes with conical apical setae; primary process present, spine-shaped, curved; adjacent row of thickened setae with globular base present on distal ¾. Tibia almost as long as femur, curved, glabrous; closing surface with a row of prostrate setae; with a patch of clavate setae apically on anterior surface. Basitarsus with long lanceolate process; proximal ½ with a patch of clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid and hind legs. Mid-leg covered with abundant fine setae, tibia unmodified, thin. Hind leg covered with fine setae, tibia unmodified, only slightly thickened. Wings. Forewing elongated and narrow, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, 7–13 subcostal veinlets, usually simple, a single forked; pterostigma rectangular, 3.5× wider than long; subcostal space with single, medially located crossvein; Sc bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight, with three or four veins arising from it, two or three from rarp1; M fused basally to R; RP base near separation of M and R, M forked slightly beyond R fork; 1r-m located between RP base and M base, forming a trapezoidal cell; 4–7 gradate crossveins present; Cu deeply forked, CuP basally angled, approaching A1, forked proximally at level of 1m-cu; A1 distally forked, A2 forked at level of CuP angle. Hind wing smaller and narrower than forewing; costal space narrow and reduced, with 4–9 veinlets; C and Sc fused at proximal 1/3 of wing length, Sc vein strongly curved posteriad at proximal margin of pterostigma to merge the RA; subcostal space without crossveins; pterostigma elongated, narrow; radial space with straight crossvein; three veins arising from rarp1, two to three from rarp2. M vein forked beyond R fork; Cu deeply forked; CuA sinuous, distally forked; CuP distally gently anteriorly curved, with three or four branches; A1 arched, A2 short, forked. Abdomen. Medially widened, keeled processes on tergites absent; tergites with abundant setation towards the abdominal apex; sternites setose.

Male genitalia (Fig. 9A–E). Tergite IX slightly narrower medially than laterally, laterally with anterior ½ sclerotized, with abundant, prominent, long, and thick setae, posterior ½ semi-membranous and depigmented. Sternites VIII and IX fused, fusion line distinct, sternite VIII with abundant prominent, long, thick setae. Sternite IX pentagonal in ventral view, with indented lateral margins, with lateral, oblique rows of long, thin setae; in lateral view curved ventrally; gonocoxites IX sinuous, long, base spatulate, curved; gradually thicker towards apex, where it is laterally curved, and equipped with 8–13 apical processes arranged as a brush. Ectoproct, anterior and posterior margins subparallel-sided, posterior ½ covered with prominent, long, and thick setae; ventrally on inner surface a sclerotized region, covered with ~ 15 stout setae, is present. Gonocoxites X thin, anterior apex spatulate, dorsally bent; posterior apex with short lateral and dorsal processes; gonostyli X thin, concave in lateral view, the rest of the structure whip-shaped, recurved. Gonapophyses X straight, narrow, anterior apex spatulate, bent dorsolaterally; gonapophyses subparallel, joined by membrane. Gonocoxites XI V-shaped in ventral view, medial lobe slightly sclerotized, rounded, bent ventrally; anteromedially sclerotized, with two plates fused to form a bridge; lateral arms of gonocoxites XI thin, straight. Hypandrium internum triangular, keeled, with lateral fins.

Figure 9. 

Anchieta fasciatellus (Westwood, 1867) A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca and gonapophyses VIII.

Female genitalia (Fig. 9F–H). Sternite VII quadrangular, with straight posterior margin; gonocoxites VIII as narrow arc-shaped sclerite; gonapophyses VIII medial part narrow, short, boat-shaped, with a blunt, dorsally curved process; lateral part of gonapophyses as a broad, trapezoidal plate, hidden under the tergite IX + ectoproct. Tergite VIII almost as wide medially as laterally, enclosing the segment VIII spiracle; lateral margin subtrapezoidal. Tergite IX + ectoproct unpaired, forming an elongated, ring-shaped sclerite, laterally wider than medially, lateral margin quadrangular. Gonocoxites IX elongated, curved, narrow, as long as the last four abdominal segments together. Gonapophyses IX as two small, setose sclerites, located behind base of gonocoxites IX. Bursa copulatrix funnel-shaped, short, narrow, membranous, striated. Spermatheca short and simple; proximal section long, thin, with a few convolutions; medial section thicker than proximal section, forming three convolutions, with transparent, digitiform diverticulum; distal section markedly wider than medial section, progressively expanded towards apex, sac-like. Fertilization canal duct long, spiral-shaped, forming four convolutions; fertilization canal short, reniform, covered with microfilaments.

Distribution

Colombia (Bolívar, Cesar, Magdalena, Santander), Panama (Canal Zone, Darien, Panamá, Panamá Oeste).

Remarks

This is a rather striking species of Anchieta previously known from Colombia and Panama (Ardila-Camacho et al. 2018). In this study, the first record for the Colombian department of Cesar is presented. In the original description by Westwood (1867), the type locality was specified as Sactam Martham, Venezuela, but the type actually has labels with “Sanctam Martham, Bolivia” as collecting information. This led to a great confusion that was clarified by Ardila-Camacho et al. (2018) and Tauber et al. (2019), who indicated that this locality actually corresponds to the city of Santa Marta (Magdalena), which is located in the Caribbean coast of Colombia.

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 Ardila-Camacho and García (2015) and Ardila-Camacho et al. (2018), this species has a mimicry pattern resembling wasps of the family Braconidae, which have defensive repugnant glands on the abdomen. Furthermore, the striking coloration with orange and black or dark brown bands or spots may serve as a defensive signal against potential predators, as these combinations of colors are typical of aposematic species.

Anchieta fumosellus (Westwood, 1867)

Figs 10, 11

Mantispa (Trichoscelia) fumosella Westwood, 1867: 504. Holotype: male, Amazonia (OUMNH), specimen examined.

Anisoptera fumosella (Westwood, 1867). Gerstaecker (1888).

Trichoscelia fumosella Westwood, 1867. Enderlein (1910).

Anchieta fumosella (Westwood, 1867). Penny (1982a).

Anchieta nobilis Navás, 1909: 484, male, female. Lectotype, sex not indicated, Brazil (MNHN). Synonymized with Anchieta fumosella by Penny (1982a): 4019, High resolution images examined.

Trichoscelia nobilis (Navás, 1909). Enderlein (1910).

Material examined

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; OUMNH.

Lectotype of Anchieta nobilis.

Brazil • ♂; Goiás, Jataí; Sep.–Nov. 1897; Anchieta nobilis Navás, Longin Navas det. 1907; Lectotype ♂ designated by R.G. Beard, 1968; MNHN.

Paralectotype of Anchieta nobilis.

Brazil • ♀; Goiás, Jataí; Anchieta nobilis Navás, Longin Navas det. 1907; Lectoallotype ♂ designated by R.G. Beard, 1968; MNHN.

Other material

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; INPA. • 1 ♂; Encruzilhada; 960 m; Nov. 1972; Seabra and Alvarenga leg.; DZUP. – Minas Gerais • 1♀; Feb. 1932; R. Bandens, J. Blaser leg.; MCZ. • 1 ♂; same data as for preceding, Passos; Mar. 1961; C. Elias leg.; det. L.A. Stange; FSCA. – Paraná • 1 ♀; São José dos Pinhais; 25°36'18"S, 49°11'37"W; 1–30 Nov. 2019; A.C. Domahovski leg.; malaise; DZUP. • 1 ♀; same data as for preceding; 17–31 Dec. 2016; A.C. Domahovski leg.; sweep; DZUP. • 1 ♂; Campo Largo, Estrada do Cerne, Km 45; 22 Nov. 1979; Expedition Dep. Zoo UFPR leg.; DZUP. • 1 ♀; Ponta Grossa, Pq. Est. Vila Velha; 25°14'S, 49°59'W; 23 Nov. 2001; G.A.R. Melo; DZUP. – Rio de Janeiro • 1 ♂; Nova Friburgo, Sans Souci; 1050 m; Nov. 2004; P. Grossi leg.; light; coleçao E. & P. Grossi; Anchieta fumosella (Westwood, 1867), det. R.J.P. Machado; INPA. • 1 ♂; Itatiaia, P. N. Itatiaia, casa pesquisador; 22°27'21"S, 44°36'30"W; 05 Dec. 2015; A.P.M. Santos and D.M. Takiya leg.; light trap; DZUP. – Santa Catarina • 1 ♀; Blumenau; F. Müller leg.; McLachlan Coll. B.M.; NHMUK, 1938-674, BMNH(E)-1241392. • 1 ♀; same data as for preceding; Blumenau, Virgil; 1987; v.d. Weele leg.; det. L.A. Stange; FSCA. • 1 ♂; Nova Teutonia; 27°11'S, 58°23'W; 300–500 m; Jan. 1971; F. Plaumann leg.; INPA. – São Paulo • 1 ♀; Jardim Botanico; 18 Dec. 1987; R.L. Jeane leg.; MCZ. • 1 ♂ 1 ♀; Same data as for preceding; Luiz Antonio, Estação Ecológica Jataí; 21°36'47"S, 47°45'04"W; 01 Oct. 2008; R. Lara leg.; light trap; DZUP. • 2 ♀; same data as for preceding; 16 Sep. 2009, Lara and team leg; Malaise; DZUP. • 2 ♂; same data as for preceding; 13 Feb. 2008; R. Lara and team leg.; light trap; DZUP. • 2 ♂; same data as for preceding; 12 Mar. 2008; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 17 Sep. 2008; R. Lara and team leg.; DZUP. • 3♂ 1♀; same data as for preceding; 01 Oct. 2008; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 29 Oct.2008; R. Lara and team leg.; DZUP. • 3 ♂; same data as for preceding; 27 Sep. 2008; R. Lara and team leg.; DZUP. • 2 ♂ 3 ♀; same data as for preceding; 28 Jan. 2009; R. Lara and team leg.; DZUP. • 5 ♂; same data as for preceding; 28 Feb. 2009; R. Lara and team leg.; DZUP. • 1 ♂ 1 ♀; same data as for preceding; 18 Mar. 2009; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 01 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♀; same data as for preceding; 09 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 15 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 29 Apr. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 27 May. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 30 Sep. 2009; R. Lara and team leg.; DZUP. • 1 ♂; same data as for preceding; 15 Oct. 2009; R. Lara and team leg.; DZUP. • 2 ♂; same data as for preceding; 29 Oct. 2009; R. Lara and team leg.; DZUP. • 1 ♀; same data as for preceding; 08 Dec. 2009; R. Lara and team leg.; DZUP. • 1 ♂; São Luiz Paraitinga, PESM nucleo St Virginia; 22°19'18"S, 45°05'43"W; 21 Nov. 2011; N.W. Perioto and team leg.; Malaise (7); DZUP.

Diagnosis

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.

Description

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. 10). Head. Mostly black, vertexal region with small pale brown mark above compound eye; with lateral rows of dark brown setae. Postgena with the inner ½ dark brown, outer ½ yellowish with dark brown suffusions. Antennal scape pale brown ventrally, dorsally dark brown, pedicel dark brown; flagellum brown to dark brown, except for ~ 8–10 yellow apical flagellomeres, some specimens with flagellum completely dark; toruli surrounded by pale brown. Clypeus with dark amber posterolateral corners; labrum with pale brown margins; mandible dark amber; maxillary palpus brown; labium black except ligula, pale brown, labial palpus brown, palpimacula pale. Thorax. Pronotum black. Episternum brown, postfurcasternum dark brown. Meso- and metanotum blackish brown, with pale brown setae; pteropleura mostly blackish brown, with orange areas on anepisterna, with brown setae. Foreleg. Coxa mostly dark reddish brown with yellowish area at apex; trochanter dark brown with pale brown ventral band. Femur mostly brown with paler areas on outer surface and small pale brown area dorsally at level of anteroventral primary process; tibia mostly dark reddish brown, with pale brown basal area, clavate setae pale; basitarsus pale amber with amber apex, second to fourth tarsomere pale brown. Mid- and hind legs. Mid-leg with brown coxa and trochanter, femur, and tibia pale brown at base and apex, the rest of the surface brown; tarsomeres completely pale brown. Hind leg with coxa, trochanter and femur brown, with interspersed pale and dark brown setae; tibia mostly pale brown with darker median area; tarsus pale brown. Wings. Forewing membrane pale amber, with darker area on distal ½ of costal field, radial space, subcostal space, area between R+M and CuP, and area adjacent to RP; with small dark amber spot posteriorly on anal region; pterostigma pale brown with pale posterior and distal margins; venation mostly brown with R+M, proximal ½ of CuA and A3 darker. Hind wing pale amber, base of subcostal space and anterior ½ of ra-rp1 slightly darker; pterostigma amber or pale brown, venation brown. Abdomen. Dark brown, with pale brown setae.

Figure 10. 

Anchieta fumosellus (Westwood, 1867) A male habitus B wings C head (yellow morph), frontal D head, frontal (dark morph) E pronotum (yellow morph), dorsal F pronotum (dark morph), dorsal G forefemur, anterior surface H same, posterior surface.

Chromatic variation (Fig. 10). Head. Mainly yellow, vertexal region with dark brown posterior transverse band and arc-shaped marking on frontal sutures; frons dark brown. Antennal scape bicolor, yellow on proximal ½, dark brown on distal ½; flagellum yellow to pale brown, setae pale to dark brown. Labrum pale brown, clypeus yellow; mandible yellow with pale amber apex; maxillary palpus amber; labium bicolor, yellow on mentum and brown on ligula, labial palpus amber. Thorax. Pronotum brown with lateral yellow stripes; episternum pale brown, with yellow margins; postfurcasternum pale brown medially, yellow on margins. Mesonotum dark brown with yellow lateral stripes; metanotum dark brown with medium yellow areas; pteropleura yellow with longitudinal brown bands, setae pale yellow. Foreleg. Coxa mostly yellow, with brown areas on basal ½; trochanter bicolor, yellow ventrally, brown dorsally. Femur mostly yellow, anterior surface with irregularly shaped brown marking extending from center to apex; posterior surface with a brown J-shaped marking on medial region; tibia yellow on basal ½, brown to dark brown on distal ½; basitarsus pale amber, remaining tarsomeres yellow. Mid and hind legs. Mid-leg mostly yellow with dark brown spots on trochanter and tibia. Hind leg with coxa bicolor, trochanter brown, femur mostly brown with yellow base and apex; tibia mainly yellow with brown posteromedian region. Abdomen. Brown, with yellow anterior and posterior margins of sclerites.

Morphology (Fig. 10). Head. Diamond-shaped in frontal view, smooth, vertexal region domed above compound eyes; coronal suture discrete; paraocular area concave. Antenna moniliform, flagellum with 43–61 flagellomeres, discoidal in shape, those of distal ½ wider, except at apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispherical, as wide as ½ of interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous to lateral and distal margins; in lateral view, posterior margin slightly raised, the rest of the surface straight; entire surface with scattered, thick setae, arising flush the pronotal surface. Postfurcasternum quadrangular. Mesonotum slightly wider than long, with a few scattered fine setae, metanotum ~ 3× as wide as long, mostly glabrous covered with microtrichia. Pteropleura covered with abundant fine and long setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, with abundant, fine and long setae; trochanter subtrapezoidal, densely setose, dorsally with a tuft of long and thick setae, anterior surface with blunt process. Femur robust, with abundant fine, long setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes with conical Stitz organs, proximally with a more developed sub-basal, spine-shaped process; adjacent row of thickened setae with globular base reduced to distal ¾ or ½; anteroventral row of processes reduced to proximal region and apex, composed of tubercle-shaped processes, with conical setae; basal primary process present, spine-shaped, curved; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae; apical region of anterior surface with a patch of clavate setae. Basitarsus with long lanceolate process; basal ½ with clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid- and hind legs. Mid- and hind leg covered with abundant fine setae; tibia unmodified, thin. Wings. Forewing oval, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, with 13–18 subcostal veinlets; pterostigma rectangular; subcostal space with single medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight with 3–6 veins arising from it, three from rarp1; M fused basally with R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork, forming a small trapezoidal cell; 6–8 gradate crossveins present; Cu deeply forked, CuA apically branched; CuP proximally angled, approaching A1, forked slightly beyond 1m-cu; A1 simple, A2 forked. Hind wing notably smaller and narrower than forewing; costal space narrow and reduced, with seven or eight subcostal veinlets; C and Sc fused at 1/3 of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma, to merge the RA; subcostal space without crossveins; pterostigma elongated, narrow; radial space with single sinuous crossvein; two or three veins arising from rarp1, 1–3 from rarp2. M vein forked at level of R fork; Cu deeply forked, CuA gently bent, distally forked, first branch forked; CuP distally gently bent anteriorly, near posterior wing margin, with two or three branches; CuP sometimes fused to posterior margin, and then diverges again forming an arch; A1 arched, A2 short, simple. Abdomen. Medially widened, tergites without keeled processes.

Male genitalia (Fig. 11A–E). Tergite IX notably narrower medially than laterally, lateral region with posteromedial tuft of long, thick setae, which surpass posterior margin of ectoproct. Sternites VIII and IX fused, fusion line moderately distinct; sternite IX posteroventrally projected, blunt, sclerotized, lateral margins concave. Gonocoxites IX thin, sinuous, as long as gonapophyses X, posterior apex curved laterally, flattened, blade-shaped, with sharp tip, sometimes slightly lanceolate, or with two or three tiny preapical processes. Ectoproct ovoid, with subparallel anterior and posterior margins; ventrally on inner surface with a more sclerotized region covered with 25–29 thick, conical setae on posterior ½ and thin, short setae on anterior ½. Gonocoxites X ventrally canaliculated, with anterior apex expanded and dorsally bent, posterior region with lateral and dorsal processes; base of gonostylus X thickened, triangular, concave in lateral view, the rest of the structure whip-shaped, short, with apical portion recurved. Gonapophyses X straight, narrow, with spatulate and dorsally recurved tips; gonapophyses joined by membrane, forming a V-shaped structure. Gonocoxites XI narrow, U-shaped, medial lobe expanded, composed of two concave plates medially joined by narrow bridge; lateral arms slightly arched, anterior apex curved.

Figure 11. 

Anchieta fumosellus (Westwood, 1867) A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 11F–I). Sternite VII subrectangular, with the posterior margin broadly concave, densely setose towards posterior margin. Tergite VIII slightly narrower medially than laterally, enclosing the spiracle of the segment, lateral margin trapezoidal. Gonocoxites VIII as two medially fused, trapezoidal, concave plates; gonapophyses VIII as a convex plate, posteromedially projected, forming a tubular process with blunt to somewhat pointed apex; lateral part of gonapophyses VIII as a trapezoidal plate hidden by the tergite IX + ectoproct. Tergite IX + ectoproct elongated, lateral margin quadrangular. Gonocoxites IX elongated, sinuous, and narrow, as long as the last five abdominal segments together. Gonapophyses IX reduced to a pair of tiny sclerites hidden by membranes, situated behind the gonocoxites IX base. Bursa copulatrix narrow, short, membranous. Spermatheca complex and coiled; proximal section long and thin, forming four coils; medial section thicker than proximal section, short, forming a couple of convolutions; distal section wider than medial section, progressively expanded towards apex, sac-like, forming a convolution; fertilization canal duct, long, spiral-shaped, with four or five convolutions, covered by the proximal section of the spermatheca; fertilization canal elongated, thin, J-shaped, covered with microfilaments.

Distribution

Brazil (Bahía, Distrito Federal, Goiás, Minas Gerais, Rio de Janeiro, Paraná, Santa Catarina, Rio Grande do Sul, São Paulo, Tocantins).

Remarks

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 (Penny 1982a; Penny and da Costa 1983; Carvalho and Corseuil 1991; Alvim et al. 2019; Schuster and Machado 2021). Herein, most of these records are confirmed based on the material examined. The holotype of Mantispa (Trichoscelia) fumosella lacks specific collecting data, but Westwood (1867) in the original description of the species indicated “Amazonia” as the collecting site. This information was questioned by Penny (1982a) and herein such idea is corroborated, as this species is restricted to Central and Southwestern Brazil. Araújo et al. (2021) reported several specimens reared from artificial nests of bees in the Amazonian portion of the Brazilian state of Mato Grosso which might be A. fumosellus, however detailed examination of such material was not possible at present, so the presence of this species in that state will require corroboration (R.J.P. Machado, pers. observation).

Anchieta fumosellus expresses a Batesian mimicry pattern with wasps of the family Vespidae (Buys 2008; Rasmussen and Ardila-Camacho 2021). Penny (1982a) realized that this species exhibits a considerable variation in the body color pattern. Some specimens can be completely dark brown, other can be yellow with dark brown stripes, while other can have intermediary colorations, tracing different species of vespid wasps as happens in the mantispid genus Climaciella Enderlein, 1910 (Rasmussen and Ardila-Camacho 2021). However, the male genitalia and wing coloration and venation between the yellow and dark forms are identical. This led to the synonym of A. nobilis (whose type has yellow body with dark stripes) with A. fumosella (in which the type has a dark brown body) (Penny 1982a). Herein that hypothesis is confirmed, after a careful examination of the morphology of several specimens of both morphs. Based on the distribution and coloration pattern and variation of this species, the model for the dark morph is likely the wasp Agelaia vicina (de Saussure, 1854), whereas the model species for the yellow morph is probably Agelaia multipicta (Haliday, 1836) (R. Lopes, pers. comm.).

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 (Peters et al. 2017).

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.

Anchieta nebulosus Ardila-Camacho & Machado, sp. nov.

Figs 12, 13

Type locality

Brazil, Espírito Santo: Linhares, Estrada, Fazenda St. Terezinha, 50 m a.s.l., Oct. 2004, P. Grossi leg.

Material examined

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)”; INPA. Paratypes. Brazil • 1 ♀; Bahía; McLachlan leg. B.M.; NHMUK, 1938-674, BMNH(E)-1241393. • 1 ♂; same data as for preceding; Camacã, Reserva Serra Bonita, Fazenda Santa Barbara; 14 Jan. 2007; A. Raw leg.; DZUP-381880.

Etymology

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.

Diagnosis

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.

Description

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. 12). Head. Mostly dark reddish brown, vertex with two dark lateral bands, alternating with paler areas, medially and posterolaterally; with rows of dark brown setae present. Postgena pale brown. Antennal scape and pedicel dark reddish brown, flagellum dark brown, except for three paler preapical flagellomeres. Mandible dark reddish brown; maxillary palpus dark amber; labium brown, palpus dark amber, except on area adjacent to articulations pale brown, palpimacula pale brown. Thorax. Pronotum dark reddish brown with a pair of pale brown markings on lateral edges, setation black. Episternum and postfurcasternum dark reddish brown. Meso- and metanotum dark reddish brown, mesonotum with black bristles. Pteropleura dark reddish brown. Foreleg. Coxa dark reddish brown; trochanter dark reddish brown. Femur mostly dark reddish brown with pale brown, medial marking, adjacent to closing surface on anterior and posterior surface; dorsal surface with a large pale brown area at apex; tibia brown, clavate setae pale brown; basitarsus brown, becoming paler towards apex; second to fourth tarsomere brown. Mid and hind legs. Mid-leg with coxa, trochanter, femur, and tibia dark brown; tarsus pale brown, with dark reddish-brown setae. Hind leg with coxa, trochanter, femur, and tibia brown, with dark brown setae; tarsus pale brown, with brown setae. Wings. Forewing membrane mainly hyaline, dark amber at base; 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; other weakly pigmented areas include posterior radial cells, area between CuP branches, and distal part of cubitoanal space; pterostigma with proximal 2/3 pale brown, distal 1/3 pale yellow; venation mostly brown, becoming paler towards posterior margin. Hind wing mostly hyaline, dark amber at base and with broad, amber, transverse band on proximal 1/3 of wing; basal ½ of subcostal space and mcu1 amber; pterostigma brown, venation brown, setation dark brown, except on paler apical margin. Abdomen. Tergites dark reddish brown except on abdominal segments IX and X + ectoproct pale brown, setation dark brown on segments I–VI, pale brown on the rest; sternites pale brown; pleural membrane dark brown.

Figure 12. 

Anchieta nebulosus Ardila-Camacho & Machado, sp. nov. A male habitus B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface G hind tibia, inner surface.

Morphology (Fig. 12). Head. Diamond-shaped in frontal view, rugose, vertex domed above compound eyes, paraocular area concave; supra-antennal region with lateral raised, elongate areas; coronal suture distinct. Antenna moniliform, flagellum slightly dorsoventrally flattened, with 47 flagellomeres, discoidal in shape, those of apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispheric, as wide as 0.5 of the interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous to lateral and distal margins; in lateral view, posterior margin slightly raised, the rest of the surface straight; entire surface with abundant long, thick setae arising flush the pronotal surface. Postfurcasternum quadrangular. Mesonotum 1.5× as wide as long, with abundant long, thick setae; metanotum ~ 3× as wide as long, mostly glabrous. Pteropleura covered with abundant long, thin setae. Foreleg. Coxa as long as femur, cylindrical, slightly distally widened, with abundant fine, long setae; trochanter subconical, densely setose, dorsally with a tuft of long, thick setae, anterior surface with conspicuous, blunt process. Femur robust, with abundant fine, long setae, longer towards ventral surface; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes, proximally with a more developed, spine-shaped, sub-basal process, adjacent row of thickened setae with globular base reduced to distal ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes, primary, spine-shaped, curved process present, adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae; anterior surface with patch of clavate setae at apex. Basitarsus with long lanceolate process; anterior surface with patch of clavate setae on basal ½, ventrally with a row of prostrate setae. Mid- and hind leg. Mid-leg with coxa, trochanter and femur with abundant fine and long setae; tibia slightly medially expanded, with abundant thick, long setae; basitarsus as long as the next three tarsomeres together; all tarsomeres covered with short, thin setae. Hind leg with coxa, trochanter and femur covered with abundant, fine, long setae; tibia notably expanded, laterally flattened, oar-shaped, with abundant long, thin setae; tarsomeres with short, thin setae, basitarsus almost as long as the next three tarsomeres together. Wings. Forewing oval, venation setose, trichosors present along the wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, 10–12 subcostal veinlets present, distal veinlets sometimes forked; pterostigma rectangular; subcostal space with single medial crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight, with three or four veins arising from it, three or four veins arising from rarp1; M fused basally to R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork forming a small, trapezoidal cell; 4–6 gradate crossveins present; Cu deeply forked, CuP basally angled, approaching A1, forked slightly beyond 1m-cu; A1 simple, A2 forked. Hind wing notably smaller and narrower than forewing, oval; costal space narrow and reduced, 6–10 veinlets; C and Sc fused at proximal 1/3 of wing length; Sc abruptly curved posteriad at proximal pterostigma margin to merge the RA; pterostigma elongated, oval; radial space with single, smoothly sinuous crossvein; two veins arising from rarp1, one or two from rarp2. M forked oppositely or slightly beyond the level of R fork. Cu vein deeply forked, CuA slightly concave, distally forked, first branch simple; CuP distally, notably, anteriorly bent, near posterior wing margin, with three or four branches; A1 arched, A2 short, simple. Abdomen. Medially widened, tergites of abdominal segments III–VI with posteromedial, keeled processes, not enlarged. Sternites covered with scattered, long, fine setae.

Male genitalia (Fig. 13A–E). Tergite IX notably narrower medially than laterally, lateral margin quadrangular, with posterior, small notch; above this notch a tuft of long, thickened, pedicellate setae, which surpass the posterior margin of ectoproct. Sternites VIII and IX, with fusion line barely perceptible; sternite VIII with abundant long, thin setae on posterior margin; sternite IX quadrangular, lateral margins with anterior slight notch; posterolateral corners rounded with short, thin setae; posterior margin ventrally curved. Gonocoxites IX short, remarkably thin, sinuous, almost as long as lateral arms of gonocoxites XI; base spatulate, rounded; apex pointed, without processes, situated under ectoprocts. Ectoproct ovoid, with short, pedicellate setae, more elongated posteroventrally; ventrally on inner surface with more sclerotized concave region, with patch of short, thickened setae. Gonoxites X forming short, triangular, thickened sclerite; posterior apex with dorsal and lateral processes; gonostyli X with triangular base, with dorsal surface concave, the rest of the structure whip-shaped, short, ventrally curved, with apical portion forming a loop. Gonapophyses X long, parallel, gently curved, with slightly dorsally recurved anterior apex; posterior apex with the surrounding membrane more sclerotized, forming a bilobed structure set with minute granules; gonapophyses joined by membrane. Gonocoxites XI arched, medial lobe composed of two quadrangular, widened plates medially joined; anterior margin with a pair of blunt, curved, projections; lateral arms of gonocoxites XI short, narrow, straight with anterior apex spatulate and rounded. Hypandrium internum arched, keeled.

Figure 13. 

Anchieta nebulosus Ardila-Camacho & Machado, sp. nov. A male terminalia B same, ventral C male genitalia, lateral D same, dorsal E same ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 13F–I). Sternite VII (gonocoxites VII) as two quadrangular, lateral plates, medially fused through a narrow bridge, with concave posterior margin. Tergite VIII slightly narrower medially than laterally, enclosing the segment spiracle, lateral margin quadrangular, notched. Gonocoxites VIII as two lateral, trapezoidal, concave plates; gonapophyses VIII forming a cubic structure, with concave sides, and with an anteromedial incision; lateral part of gonapophyses VIII as a trapezoidal, elongated plate, hidden under tergite IX + ectoproct. Tergite IX + ectoproct triangular. Gonocoxites IX narrow, elongated, as long as the last four abdominal segments. Gonapophyses IX reduced to a pair of tiny sclerites hidden by tergite IX + ectoproct. Bursa copulatrix funnel-shaped membranous, short. Spermatheca tightly coiled; proximal section long, thin, forming three coils on distal part; medial section, short, thicker than proximal section, forming few convolutions; distal section remarkably wider than medial section, strongly expanded towards apex, sac-shaped; fertilization canal duct long, spiral-shaped, with five convolutions, with widened at preapical part; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Bahía, Espírito Santo).

Remarks

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 (Rasmussen and Ardila-Camacho 2021).

Anchieta nothus (Erichson, 1839)

Figs 14, 15

Mantispa notha Erichson, 1839: 170. Holotype: male, Brazil (ZMB), specimen examined.

Trichoscelia notha (Erichson, 1839). Hagen (1861).

Symphrasis notha (Erichson, 1839). Navás (1909).

Anisoptera notha (Erichson, 1839). Enderlein (1910).

Anchieta notha (Erichson, 1839). Penny (1982b).

Material examined

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]; ZMB 123.

Other material

Brazil • 1♀; Bahía; Hagen, Winthom leg.; MCZ. – Minas Gerais • 1 ♂; II.1941, Lopes & Gomes leg., det. L.A. Stange; FSCA.

Diagnosis

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.

Description

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. 14). Head. Mostly dark reddish brown, vertexal region with lateral rows of dark brown setae, supra-antennal region with small yellowish mark above toruli. Postgene yellowish with dark brown spots. Antenna dark reddish brown, except for four pale brown preapical flagellomeres. Mandible pale amber; maxillary palpus amber; labium black except pale brown ligula, labial palpus dark amber, palpimacula amber. Thorax. Pronotum dark reddish brown with yellowish markings on lateral edges; episternum and postfurcasternum dark reddish brown. Meso- and metanotum dark brown, metanotum with dark reddish brown bristles; pteropleura dark reddish brown. Foreleg. Coxa dark reddish brown; trochanter dark brown. Femur mostly dark brown with pale brown medial and apical areas. Tibia mostly brown, clavate setae pale brown; basitarsus amber with dark apex, second to fourth tarsomere pale brown tarsomere. Mid- and hind leg. Mid-leg with coxa, trochanter, femur, and tibia dark reddish brown; tarsomeres yellow with dark reddish brown setae. Hind leg with coxa, trochanter and femur dark brown; tibia with brown base and apex, medial region yellow, with concolorous setae; tarsomeres yellow with interspersed yellow and dark brown setae. Wings. Forewing membrane mostly hyaline, with pale amber irregular areas at wing base, area around the 1r-m, proximal 1/4 of wing, distal subcostal veinlets, apical area of subcostal field, and area surrounding 1ra-rp; pterostigma with proximal ¾ pale brown, distal ¼ pale yellow; venation mostly brown. Hind wing mostly hyaline with broad transverse pale amber band on proximal 1/3; costal field, basal ½ of subcostal space, and mcua1 pale amber; pterostigma brown, venation brown. Abdomen. Tergite of abdominal segments III and IX mostly yellow, the rest mostly dark reddish brown; sternites mainly brown, with yellow suffusions; pleural membrane dark brown.

Figure 14. 

Anchieta nothus (Erichson, 1830) A female habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F hind tibia, outer surface.

Morphology (Fig. 14). Head. Diamond-shaped in front view, rugose; vertexal region domed above compound eyes; coronal suture distinct; paraocular area concave; supra-antennal region with lateral, elongated, raised areas. Antenna moniliform, with 51 flagellomeres, discoidal in shape, those at the apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispheric, as wide as ¾ of interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous lateral and distal margins; in lateral view, posterior margin slightly raised; entire surface with abundant, thick setae arising flush the pronotal surface. Postfurcasternum quadrangular. Mesonotum wider than long, with abundant long and thick setae; metanotum ~ 3× as wide as long, mostly glabrous covered. Pteropleura covered with abundant fine and long setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, with abundant fine and long setae; trochanter subtrapezoidal, densely setose, dorsally with a tuft of long, thick setae, inner surface with blunt process. Femur robust, with abundant, long, fine setae; closing surface with posterolateral row of integumentary specializations fully-developed, composed of tubercle-shaped processes, proximally with a more developed, spine-shaped, sub-basal process, adjacent row of thickened setae with globular base reduced to distal ¾ or ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes; primary, spine-shaped process present, curved; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae; anterior surface with a patch of clavate setae at apex; basitarsus with long lanceolate process; proximal ½ with patch of clavate setae on anterior surface, and single row of prostrate setae ventrally. Mid- and hind leg. Mid-leg with coxa, trochanter and femur densely covered with fine and long setae; tibia moderately, medially expanded, with abundant long, thick setae; basitarsus as long as the following three together, covered with shorter setae compared to other segments. Hind leg with coxa, trochanter and femur covered with abundant, thin, long setae; tibia conspicuously expanded, laterally flattened, oar-shaped, densely setose, with setae shorter than those of mid tibia; tarsomeres with short, thin setae, basitarsus as long as the following three tarsomeres together. Wings. Forewing oval, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, 11–13 subcostal veinlets present; pterostigma rectangular; subcostal space with single, medial crossvein; Sc abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight with three or four veins arising from it, three from rarp1; M basally fused with R; RP base located near separation of M from R, M fork near such separation; 1r-m situated between RP base and M fork, forming a small trapezoidal cell; 4–6 gradate crossveins present; Cu deeply forked, CuP basally angled, approaching A1, forked slightly beyond 1m-cu; A1 simple. Hind wing notably smaller and narrower than fore wing, oval; costal space narrow and reduced, with 7–9 veinlets; C and Sc fused at 1/3 of wing length; subcostal space without crossveins; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, oval; radial space with single crossvein, two veins arising from rarp1, one or two from rarp2. M vein forked slightly beyond R fork. Cu vein deeply forked, CuA straight to slightly concave, distally simple or forked, first branch simple or forked; CuP distally, strongly, anteriorly arched, pectinate; A1 arched, A2 short, simple. Abdomen. Medially widened, tergites of abdominal segments III–VII with prominent posteromedial keeled processes; processes on tergites IV–VI notably enlarged in both sexes, in male the largest situated on tergite IV, and on tergite V in female.

Male genitalia (Fig. 15A–E). Tergite IX notably narrower medially than laterally, lateral margin rounded, posteriorly with long and thin setae. Sternites VIII and IX fused, fusion line barely perceptible; sternite VIII with abundant long, thin setae; sternite IX quadrangular, posteromedially produced into a blunt, elongated lobe; lateral margins concave, median region with curved, transverse rows of long, thin setae. Gonocoxites IX short and narrow, as long gonostyli X, sinuous, posterior apex ventrally curved, with three short apical processes. Ectoproct elongated and narrow, with subparallel anterior and posterior margins, posteroventrally with short, pedicellate setae; ventrally on inner surface with a more sclerotized region set with ~ 15 short and thick setae on posterior ½ and some fine and short setae on anterior ½. Gonocoxites X constrained at anterior 1/3, base spatulate and bent dorsally; apex with dorsal and lateral processes; base of gonostyli X triangular, ventrally concave; the rest of the structure whip-shaped, short, with apical portion recurved. Gonapophyses X slightly thickened, gently curved, anterior apex spatulate, dorsally recurved; both joined by membranes forming a V-shaped structure. Gonocoxites XI narrow, arched, medial lobe expanded, composed of two oval, widened, concave plates, joined medially by narrow bridge. Lateral arms of gonocoxites XI with short and slender.

Figure 15. 

Anchieta nothus (Erichson, 1830) A male terminalia B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca.

Female genitalia (Fig. 15F–H). Sternite VII as two lateral, broad, subtrapezoidal plates (gonocoxites VII), posteromedially fused through thin bridge, posterior margin broadly concave. Tergite VIII slightly narrower medially than laterally, enclosing the spiracle of the segment, lateral margin quadrangular, with a notch. Gonocoxites VIII as two lateral, trapezoidal, concave plates; gonapophyses VIII medial part forming a cubic structure, with concave sides, and with an anteromedial incision; lateral part a trapezoidal, elongated plate, hidden under the tergite IX + ectoproct. Tergite IX + ectoproct triangular. Gonocoxites IX narrow, elongated, as long as the last four abdominal segments. Gonapophyses IX reduced to a pair of tiny sclerites hidden by tergite IX + ectoproct. Bursa copulatrix funnel-shaped, membranous, short. Spermatheca tightly coiled; proximal section long and thin, forming three coils distally; medial section, short, thicker than proximal section, forming a pair of convolutions; distal section markedly wider than medial section, progressively expanded towards apex, sac-like; fertilization canal duct long, spiral-shaped, with five convolutions, widened at preapical part; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Bahía, Minas Gerais).

Remarks

Prior to this work, this species was known from Brazil, although information about its distribution within the country is not available (Machado 2022). The type specimen of Mantispa notha deposited in Museum für Naturkunde der Humboldt-Universität (ZMB) has only “Brasilien” as collecting locality. Penny and da Costa (1983) redescribed this species based on a female lacking collecting data. Herein the first specific records of this species in Brazil (i.e., Bahía and Minas Gerais) are presented.

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 (Rasmussen and Ardila-Camacho 2021).

Anchieta partheniellus (Westwood, 1867)

Figs 16, 17

Mantispa (Trichoscelia) partheniella Westwood, 1867: 501. Lectotype male, Paralectotype female, Amazonia (OUMNH), specimens examined.

Trichoscelia partheniella (Westwood, 1867). Enderlein (1910).

Anchieta partheniella (Westwood, 1867). Penny (1982a), Ohl (2004).

Material examined

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; OUMNH.

Other material

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; INPA. – Pará • 1 ♀; Tucurú, Río Tocantins; 03 Jun. 1984; Saúde leg.; Malaise trap; Anchieta bella (Westwood, 1867) det. R.J.P. Machado; INPA. – Rondônia • 1 ♀; 62 Km SW Ariquemes, Fazenda Rancho Grande; 11 Oct. 1993; C.W. and L.B. O’Brien leg.; CAS. • 1 ♀; same data as for preceding; UV merc. Vapor light; Anchieta bella, det. Penny, 2001; CAS. • 1 ♀; same data as for preceding; 9 Nov.1994; CAS.

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; NHMUK.

Suriname • 1 ♀; Republiek; 13 Nov. 1961; P.H.v. Doesburg leg.; A. romani, det. Geisjkes 1970; FSCA.

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.

Diagnosis

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.

Description

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. 16). Head. Mostly orange; vertexal region posteriorly with a broad transverse brown band, laterally with rows of dark brown setae; supra-antennal area with dark brown, lateral trapezoidal markings, such markings connected to the frontal markings through interantennal area. Antennal scape ventrally orange, dorsally brown, pedicel brown, flagellum brown, except for four or five yellow preapical flagellomeres. Frons with lateral brown triangular markings forming an inverted, V-shaped pattern. Clypeus with brown posterolateral corners; Labrum with brown medial suffusions; mandible orange with dark amber apex; maxillary palpus brown; labium orange, palpus brown, palpimacula pale brown. Thorax. Pronotum dark brown, with orange, transverse band on anterior 1/3, anterior margin brown; episternum bicolor, yellow on anterior ¼, dark brown on the rest; postfurcasternum pale to dark brown. Mesonotum orange with brown anterior margin, medial area of scutum with brown suffusions, posterior margin of scutellum dark brown, setae pale brown; metanotum with brown lateral markings, posterior margin of scutellum brown. Pteropleura mainly orange with some brown areas, setae orange. Foreleg. Coxa mostly orange with extensive dark brown area on posterior surface, apex slightly infuscated; anterior surface with interspersed orange and dark brown setae; trochanter orange except dorsally brown. Femur orange, with enlarged dorsal spot extending towards anterior surface; posterior surface with brown preapical longitudinal mark, setation mostly orange, except on medial region dark brown; tibia bicolor with orange basal ½, brown distal ½; basitarsus pale brown with amber apex, remaining tarsomeres orange. Mid- and hind leg. Mid-leg predominantly orange, with brown suffusions. Hind leg orange; femur with brown medial suffusion; tibia with posterodistal margin and distal 1/3 of anterior surface black. Wings. Forewing membrane hyaline, with yellow areas at wing base and area adjacent to proximal region of pterostigma; a yellow transverse band R fork level is present; pterostigma bicolor, orange on proximal ½, yellow on distal ½; venation predominantly orange with dark brown suffusions at base of RA and jugal lobe. Hind wing hyaline, base of subcostal space orange; pterostigma orange with black suffusions; venation orange. Abdomen. Tergite I yellow with brown area at center, tergite II with lateral brown markings, tergites III and IV yellow with two brown, lateral bands, tergites V–VIII brown; sternites yellow.

Figure 16. 

Anchieta partheniellus (Westwood, 1867) A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface G hind tibia, outer surface H same, inner surface.

Morphology (Fig. 16). Head. Diamond-shaped in frontal view, smooth; vertexal region domed above compound eyes, laterally with rows of short, reclined setae; paraocular area concave; coronal suture discrete. Antenna moniliform, flagellum with 51 flagellomeres, discoidal in shape, except for those of apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispherical, as wide as 1/2 of interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous lateral and distal margins; posterior region slightly raised; entire surface with scattered, thick setae arising flush the pronotal surface. Episternum with long, thin setae; postfurcasternum quadrangular. Mesonotum slightly wider than long, with few long, fine setae; metanotum ~ 3× as wide as long, glabrous. Pteropleura covered with abundant fine, long setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, with abundant fine, long setae; trochanter semi-triangular, densely setose, dorsally with tuft of long, thick setae, anterior surface with blunt process. Femur robust, covered with abundant long, fine setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes, proximally with sub-basal, spine-shaped process, adjacent row of thickened setae with globular base present on distal ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes, with conical setae; primary, spine-shaped, basal process present, curved; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, with a row of prostrate setae on closing surface, anterior surface with apical patch of clavate setae. Basitarsus with long lanceolate process; anterior surface with patch of clavate setae on proximal ½, ventrally with single row of prostrate setae. Mid- and hind leg. Mid-leg covered with abundant, fine, long setae; tibia slightly widened. Hind leg densely covered with long, fine setae, except on tarsus shorter; tibia conspicuously widened and laterally flattened, oar-shaped. Wings. Forewing oval, venation densely setose, trichosoros present along wing margin except at base; costal space slightly widened medially, humeral vein forked, 10–12 subcostal veinlets present; pterostigma rectangular; subcostal space with single, medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight with four veins arising from it, four from rarp1; M fused basally to R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork, forming a small trapezoidal cell; six gradate crossveins present; Cu deeply forked, CuA distally branched, CuP proximally angled, forked; A1 simple. Hind wing oval, notably smaller and narrower than forewing; costal space narrow and reduced, with seven or eight subcostal veinlets; C and Sc fused at 1/3 of wing length, subcostal space without crossveins, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, slightly widened distally; radial space widened, with single, sinuous crossveins; two veins arising from rarp1, two or three from rarp2. M forked at R fork level. Cubitoanal space with distal crossveins. Cu vein deeply forked, CuA gently curved to nearly straight, distally forked, first branch simple; CuP distally anteriorly bent near wing margin, with three branches; A1 arched, A2 short, simple. Abdomen. Medially widened, without keeled processes, tergites V–IX with abundant long, thin setae; sternites densely setose.

Male genitalia (Fig. 17A–E). Tergite IX medially narrower than laterally, posterior corner with abundant long, thick setae. Sternites VIII and IX fused, fusion line weakly marked; sternite IX anteriorly with long, thick setae, medially with short setae, distal ½ glabrous with concave lateral margins; anterior ½ somewhat constricted. Gonocoxites IX short, base spatulate; apex strongly recurved lateroventrally, with two apical and four preapical processes on outer surface. Ectoproct ovoid, posteroventrally with patch of 22 or 23 thickened, dark setae; outer surface with smooth, glabrous, bulging area; inner surface with sclerotized ventral area, with a patch of microtrichia. Gonocoxites X medially constricted, hourglass-shaped; anterior ½ thickened, expanded, dorsally bent, ventrally canaliculated; posterior ½ triangular, with paired dorsal and lateral processes; gonostyli X base triangular, thickened, with curved lateral processes, dorsally concave, the rest of the structure whip-shaped, short, apical portion recurved. Gonapophyses X rod-shaped, straight, narrow, anterior apex spatulate, dorsally curved, posterior apex dorsally curved, surrounding membrane with minute granules; gonapophyses joined by membrane, forming a V-shaped structure. Gonocoxites XI thin, U-shaped, medial lobe flattened, weakly sclerotized at center, posterior margin almost straight, lateral margins emarginate. Lateral arms of gonocoxites XI straight, with anterior apex curved.

Figure 17. 

Anchieta partheniellus (Westwood, 1867) A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 17F–I). Sternum VII (gonocoxites VII) composed of two medially fused trapezoidal lateral plates, posterior margin obtuse. Tergite VIII slightly narrower medially than laterally, enclosing spiracle of the segment, lateral margin rounded. Gonocoxites VIII as two concave, medially fused, trapezoidal plates; gonapophyses VIII medial part ventrally projected forming a canal, with medial depression on ventral surface, distal margin rounded, ventrally curved; lateral part of gonapophyses VIII a trapezoidal plate, forming a chamber-shaped plate, dorsally with two convex areas, located behind the genital pore. Tergite IX + ectoproct triangular. Gonocoxite IX long, straight, narrow, as long as the last five abdominal segments together. Bursa copulatrix unsclerotized, long, conical; spermatheca spiral-shaped; proximal section long, thin, progressively thickened, spiral-shaped, forming three coils; medial section slightly narrower than thickest part of proximal section, entangled, forming several convolutions; distal section thicker than proximal and medial sections, progressively wider towards the apex. Fertilization canal duct long, spiral-shaped, forming two coils; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Amazonas, Pará, Rondônia), Colombia (Meta), Ecuador (Napo), Suriname, Venezuela (Amazonas).

Remarks

Lectotypes (male and female) of this species deposited in the Oxford University Museum have only “Amazons” as collecting data. Penny (1982a) and Penny and da Costa (1983) redescribed this species, but no additional records were presented. Although the types of this species were likely collected in Brazil, so far there are no specific state records of this species in the country. Herein, the first records for the states of Amazonas, Pará, and Rondônia (all in the Amazon region) are presented, and the first records from Colombia, Ecuador, Suriname, and Venezuela are provided.

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 (Rasmussen and Ardila-Camacho 2021). Considering the relationships between these species, this coloration pattern apparently arose several times within this group of species of Anchieta.

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.

Anchieta remipes (Gerstaecker, 1888)

Figs 18, 19

Anisoptera remipes Gerstaecker, 1888: 120. Holotype: male, Colombia, Bogotá (ZIMG), high resolution images examined.

Trichoscelia remipes (Gerstaecker, 1888). Penny (1977), Hoffman (2002), Ohl (2004).

Anchieta remipes (Gerstaecker, 1888). Ardila-Camacho and García (2015).

Material examined

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; ZIMG.

Diagnosis

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.

Description

Measurements. Male (n = 1). Forewing length: 8.9 mm; Hind wing length: 5.0 mm.

Coloration (Fig. 18). Head. Mostly orange, vertexal region with posterior, transverse brown band; supraantennal region with lateral triangular marks; with lateral rows of brown setae. Antennal scape orange. Frons with brown suffusions. Clypeus orange laterally, brown at center; labrum medially brown, orange on edges; mandible orange with amber apex; maxillary palpus orange; labium orange, palpimacula orange. Thorax. Pronotum dark brown with orange, transverse band on anterior 1/3, anterior margin brown; episternum pale brown with darker margins, postfurcasternum dark brown with poster paler area. Mesonotum orange with brown areas; metanotum mostly brown. Pteropleura with episterna orange with brown areas, epimera brown. Foreleg. Coxa brown, with distal 1/3 orange, setation brown; trochanter orange except dorsally brown. Femur orange with pale brown, irregular areas on posterior and dorsal surfaces; anterior surface with medial area pale brown; setation mostly orange; basal ½ of tibia orange, distal ½ brown; clavate setae orange; tarsus orange. Mid- and hind leg. Mid-leg mostly orange. Hind leg mostly orange; tibia with posterior margin and preapical region with brown suffusions. Wings. Forewing membrane hyaline; pterostigma bicolor, pale brown on proximal ⅔, pale yellow on the rest; venation with orange and brown areas. Hind wing hyaline, pterostigma pale brown, venation mostly brown, most of C and Sc orange. Abdomen. cleared.

Figure 18. 

Anchieta remipes (Gerstaecker, 1888) A male habitus, dorsal B wings C head, frontal D forefemur, anterior surface E head and thorax, lateral F hind tibia, outer surface.

Morphology (Fig. 18). Head. Diamond-shaped in frontal view; vertexal region domed above compound eyes, smooth, laterally with rows of short, reclined setae; paraocular area concave; coronal suture discrete. Antennal scape 2× as long as wide. Compound eye hemispherical, as wide as ½ the interocular distance at toruli level. Thorax. Pronotum as long as wide, with groove contiguous to lateral and distal margins; posterior region slightly raised, entire surface with abundant, thick setae arising flush the pronotal surface; episternum with long, thin setae; postfurcasternum trapezoidal. Mesonotum 1.5× as wide as long, with few long, thickened setae; metanotum ~ 3× as wide as long, glabrous. Pteropleura covered with abundant long, thin setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, densely covered with abundant fine, long setae; trochanter conical, densely setose, dorsally with tuft of long, thick setae, anterior surface with blunt process. Femur robust, densely set with fine setae; closing surface with posteroventral row of integumentary specializations fully-developed, composed of tubercle-shaped processes, proximally with spine-shaped, sub-basal process, adjacent row of thickened setae with globular base present on distal ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes; primary process present, curved; adjacent row of thickened setae with globular base present on distal ⅘. Tibia almost as long as femur, curved, glabrous, with a row of setae prostrate on closing surface, and a patch of clavate setae apically on anterior surface. Basitarsus with lanceolate process reaching middle of fourth tarsomere, proximal ½ with patch of clavate setae on anterior surface, ventrally with single row of prostrate setae. Mid- and hind leg. Mid-leg covered with abundant fine, long setae, tibia slightly widened. Hind leg densely covered with long, fine setae, except on the tarsi, shorter, tibia notably widened, laterally flattened, oar-shaped. Wings. Forewing oval, venation densely setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein forked, 11–13 subcostal veinlets present; pterostigma rectangular; subcostal space with single, medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight, with three veins arising from it, two from rarp1; M basally fused to R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork forming a small trapezoidal cell; three gradate crossveins present; CuA distally branched; CuP angled proximally, touching A1, forked nearly at M fork level; A1 simple, A2 forked. Hind wing oval, notably smaller and narrower than forewing; costal space narrow and reduced, with five veinlets; C and Sc fused at proximal 1/3 of wing length, subcostal space without crossveins; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, slightly widened distally; radial space widened with single reclined crossvein; one or two veins arising from rarp1, 0–2 from rarp2. M vein forked slightly beyond R fork. Cu vein deeply forked, CuA slightly bent at mid-length, distally forked; CuP distally bent anteriorly near posterior wing margin, with three branches; A1 arched, A2 short, simple.

Abdomen. Cylindrical, without keeled processes.

Male genitalia (Fig. 19). Tergite IX medially narrower than laterally, laterally with abundant long, thin setae. Sternites VIII and IX fused, fusion line weakly marked; sternite VIII rectangular, with long, thickened setae; sternite IX trapezoidal, with broad posteromedial, quadrangular lobe. Gonocoxites IX fusiform, reduced. Ectoproct triangular, ventrolaterally with smooth, glabrous, bulging area. Gonocoxites X forming an elongate sclerite; anterior dorsally curved; posterior apex expanded; base of gonostyli X triangular, thickened, with lateral processes; the rest of the structure whip-like, short, dorsally recurved, and anteroventrally projected. Gonapophyses X elongate, narrow; gonapophyses joined by membrane, forming a V-shaped structure, posterior apex of both fused, forming a blunt process. Gonocoxites XI thin, U-shaped, medial lobe flattened, posterior margin rounded; lateral arms of gonocoxites XI straight.

Figure 19. 

Anchieta remipes (Gerstaecker, 1888) A male terminalia, lateral B same, ventral.

Distribution

Colombia (Cundinamarca?).

Remarks

This species is so far known only from the original description by Gerstaecker (1888). In this description the only collecting data available is “Bogotá”, the main city of Colombia, located in the Cundinamarca department. However, the labels of the holotype only have “Columb.” As collecting information, so presently the distribution of the species in Colombia is unknown.

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.

Anchieta romani (Esben-Petersen, 1917)

Figs 20, 21

Anisoptera romani Esben-Petersen, 1917: 14. Holotype: male, Brazil, Rio Autaz (NHRS), specimen examined.

Trichoscelia romani (Esben-Petersen, 1917). Penny (1977).

(non) Anchieta bella (Westwood, 1867). Penny (1982a).

Material examined

Holotype. Brazil • ♂; Amazonas, Río Autaz; Sept.; Roman leg.; “Anisoptera romani, male, det. Esben-Petersen (NHRS)”, “Type red label, Rikmuseum Stockholm”, “198, 80”, “190, 57”, “NHRS-KAJO 000000370”; terminalia cleared and stored in a microvial; NHRS.

Other material

Peru • 1 ♀ 2 ♂; San Martín, Tarapoto Urku, nr. Boca Toma; 400 m. a.s.l.; 10 Jul. 2012; C. Rasmussen leg.; nest collected; ZMB. • 1 pupa; same data as for preceding; 6 Sep. 2012; ZMB.

Diagnosis

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.

Description

Measurements. Male (n = 2). Forewing length: 9.0–9.2 mm; Hind wing length: 5.2–5.4 mm.

Coloration (Fig. 18). Head. Mostly orange; vertexal region brown with triangular, yellow areas adjacent to frontal sutures; laterally with rows of dark brown setae; supra-antennal region brown with oval yellow, medial area. Antennal scape orange ventrally, dorsally pale brown on distal ½, pedicel pale brown, flagellum pale brown, except for three or four yellow, preapical flagellomeres. Frons with lateral brown triangular markings, forming an inverted V-shaped pattern. Labrum yellow, clypeus with brown suffusions at center; mandible orange with pale amber apex; maxillary palpus orange with pale brown suffusions; labium orange, palpus with pale brown suffusions, palpimacula orange. Thorax. Pronotum dark brown, except anterior 1/3 with orange transverse band; episternum brown with yellow anterior margin; postfurcasternum brown. Mesonotum orange with dark brown anterior margin, medial area with broad, brown band, scutellum with brown spot on anterior ½ and brown posterior margin; metanotum yellow with brown lateral markings, and brown medial band and anterior and posterior margins; pteropleura orange with broad brown areas, pale brown or orange setae present. Foreleg. Coxa mostly orange, with extensive pale brown spot on posterior surface, base and apex brown; trochanter orange on ventral surface, pale brown on dorsal surface. Femur orange with broad dorsal spot extending to most of anterior surface; posterior surface with preapical, longitudinal mark and brown transverse medial mark; tibia bicolor, basal ½ orange, distal ½ pale brown; basitarsus orange on basal 1/2, pale brown on distal 1/2, remaining tarsomeres orange. Mid- and hind leg. Mid-leg predominantly orange, coxa with basal 1/3 and apex pale brown, trochanter orange with pale brown suffusions, femur medially pale brown; tibia with apical 1/3 pale brown. Hind leg mainly orange, coxa with pale brown suffusions; tibia orange, except at middle of outer surface and distal ¾ of inner surface brown, tip orange. Wings. Forewing membrane hyaline, with yellow areas at base, area surrounding 1sc-r, 1ra-rp, and area adjacent to proximal region of pterostigma; pterostigma bicolor, orange on proximal ½, pale yellow on distal ½; venation predominantly orange, except wing base, medial region of R + M, subcostal veinlets, 1c-sc, 1ra-rp, 1r-m, and 1m-cu veins brown. Hind wing hyaline, base of subcostal space orange; pterostigma orange with pale brown suffusions, setation blackish; venation mainly pale brown, yellow near wing margin, wing base, and apex of C + Sc. Abdomen. Tergites I–III yellow with pale brown lateral markings, remaining tergites yellow with brown suffusions; sternites yellow; pleural membrane pale brown.

Figure 20. 

Anchieta romani (Esben-Petersen, 1817) A male habitus, dorsal (abdomen removed) B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface G hind tibia, outer surface H same, inner surface.

Morphology (Fig. 20). Head. Diamond-shaped in frontal view, smooth; vertexal region domed over compound eyes, laterally with rows of short, reclined setae; paraocular area concave; coronal suture discrete. Antenna moniliform, flagellum with 44–46 flagellomeres, discoidal in shape, except for those of apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at toruli level. Thorax. Pronotum nearly as long as wide, with groove contiguous to lateral and distal margins; posterior margin with slight outgrowth; entire surface with abundant, thick setae arising flush the pronotal surface; episternum with interspersed long, thin and short, thickened setae; postfurcasternum quadrangular. Mesonotum wider than long, with abundant long, thickened setae present on medial area; metanotum ~ 3× as wide as long, glabrous. Pteropleura covered with abundant long, thin setae. Foreleg. Coxa as long as femur, cylindrical, slightly expanded at preapical region, densely covered with fine, long setae; trochanter semi-triangular, densely setose, dorsally with tuft of long, thick setae, anterior surface with blunt process. Femur robust, densely set with long, fine setae; closing surface with posteroventral row of integumentary specializations fully-developed, composed of tubercle-shaped specializations, proximally with spine-shaped, sub-basal process; adjacent row of thickened setae with globular base present on distal ½; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped specializations; primary process present, curved; adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, with a row of prostrate setae on closing surface, and patch of clavate setae apically on anterior surface. Basitarsus with long lanceolate process, with clavate setae on proximal region of anterior surface, ventrally with single row of prostrate setae. Mid- and hind leg. Mid-leg covered with abundant fine, long setae, tibia slightly widened. Hind leg densely covered with long, fine setae, except on tarsus, shorter; tibia notably widened and laterally flattened, oar-shaped. Wings. Forewing oval, venation densely setose, trichosors present along wing margin; costal space slightly widened medially, humeral vein simple or forked, 10–12 subcostal veinlets present; pterostigma rectangular; subcostal space with single medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight, with four or five veins arising from it, three or four from rarp1; M fused basally to R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork forming a small, trapezoidal cell; five or six gradate crossveins present; Cu deeply forked, CuA distally branched; CuP proximally angled, approaching A1, forked; A1 simple, A2 medially forked. Hind wing oval, notably smaller and narrower than forewing; costal space narrow and reduced, with seven or eight veinlets; C and Sc fused at proximal 1/3 of wing length, subcostal space without crossveins; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, slightly widened distally; radial space widened with single, straight, crossvein; two or three veins arising from rarp1, 1–3 from rarp2. M vein forked at R fork level. Cu vein deeply forked, CuA slightly concave, distally forked, first branch simple; CuP distally curved anteriorly near wing margin, with 2–4 branches; A1 arched, A2 short, simple. Abdomen. Cylindrical, tergites III and IV with short, posteromedial keeled processes in both sexes, tergites V–IX with abundant long, thin setae; sternites densely setose.

Male genitalia (Fig. 21A–E). Tergite IX medially narrower than laterally, posterolaterally with abundant long, thick setae. Sternites VIII and IX fused, fusion line weakly marked; sternite VIII rectangular with abundant long, thickened setae, sternite IX U-shaped, with long, thick setae on anterior ½, posterior ½ glabrous, medially convex, forming a canal, laterally concave. Gonocoxites IX short, sinuous, with base flattened; apex laterally recurved, with two or three longitudinally arranged, preapical processes on outer surface. Ectoproct ovoid, narrow, posteroventrally with patch of 14–16 thick, dark setae; laterally with smooth, glabrous, bulging area; inner area sclerotized, with patch of short, fine setae. Gonocoxites X forming a triangular sclerite; anterior apex laterally flattened, ventrally concave; posterior apex with paired dorsal and lateral processes; base of gonostyli X U-shaped, thickened, concave, set with curved, lateral processes; the rest of the structure whip-like, short, recurved, apical portion forming a loop. Gonapophyses X straight, narrow, anterior apex spatulate, dorsally curved, posterior apex dorsally curved, surrounding membrane set with minute granules; gonapophyses joined by membrane, forming a V-shaped structure. Gonocoxites XI thin, U-shaped, medial lobe flattened, quadrangular, weakly sclerotized at center, anterior margin with U-shaped concavity; lateral arms of gonocoxites XI gently sinuous, anterior apex curved.

Figure 21. 

Anchieta romani (Esben-Petersen, 1817) A male terminalia, lateral B same, ventral C male genitalia, lateral D same dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 21F–I). Sternite VII (gonocoxites VII) formed by two medially fused trapezoidal lateral plates, covered with long and thin setae, posterior margin forming an obtuse angle. Tergite VIII almost as wide medially than laterally, laterally enclosing the spiracle of the segment, lateral margin quadrangular. Gonocoxites VIII forming a narrow trapezoidal plate, with two lateral concavities; gonapophyses VIII medial part forming an enlarged canal, with a depression on ventral surface, apex ventrally curved, rounded; lateral part of gonapophyses VIII forming a chamber-shaped dorsal structure with two lateral concavities. Tergite IX + ectoproct triangular. Gonocoxite IX long, straight, narrow, as long as the last five abdominal segments together. Gonapophyses IX as a pair of tiny sclerites hidden under tergite IX + ectoproct. Bursa copulatrix unsclerotized, long, progressively narrower; spermatheca complex and coiled; proximal section thin, slightly thickened at coiled area, forming three convolutions; medial section thicker than proximal section, long, coiled, forming multiple convolutions; distal section wider than medial section, sac-shaped, forming a loop. Fertilization canal duct long, spiral-shaped, forming three convolutions, slightly preapically expanded; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Brazil (Amazonas), Peru (San Martín).

Remarks

This species was synonymized with A. bellus by Penny (1982a) without examination of the type of Anisoptera romani Esben-Petersen, 1917. Both species have basically the same coloration pattern, but the hind tibia of A. bellus is narrower than that of A. romani, while the shape of the male gonocoxites IX and X are noticeably different. Based on these characters, herein Anchieta romani (Esben-Petersen, 1917) is reinstated as a valid species.

Regarding its distribution, the collecting data of the type of A. romani appears as “Rio Autaz, Amazon”, locality that was corrected by Penny (1982a) as Autazes, in the Brazilian state of Amazonas. Here, this species is reported from Peru (San Martin) for the first time.

In a recent publication by Rasmussen and Ardila-Camacho (2021) this species was referred as Anchieta sp. nov. Since the taxonomic status of A. romani was not clarified at that time, the specimens cited in that publication were initially interpreted as belonging to a new species. In that contribution, aspects of the life history of this species like its association with the vespid wasp Montezumia dimidiata Saussure, 1852 (Vespidae: Eumeninae) and its mimicry with Ptilotrigona lurida (Smith, 1854) were discussed.

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.

Anchieta sophiae Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 22, 23

Type locality

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.

Material examined

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” MNHN. Paratypes. French Guiana • 1 ♂; same data as for holotype; CNIN. – Maripasoula • 1 ♀; Contreforts du Mitaraka, crique Alama, Fôret vallonnée au pied d’inselbergs; 12 Mar. 2015; light trap; CSCA.

Diagnosis

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.

Etymology

First author dedicates this species after his daughter Lauren Sophia Ardila, born on 29 of December of 2015.

Description

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. 22). Head. Mostly yellow, vertexal region with transversal, dark brown band; with lateral rows of dark brown setae; supra-antennal area with lateral, triangular, dark brown marks. Antennal scape yellow with brown apex, pedicel brown; flagellum pale brown, except apex yellow. Frons with central, dark brown dot; mandible pale brown, with base yellow; maxilla with galea and maxillary palpus pale brown; labium yellow, labial palpus pale brown, palpimacula yellow. Thorax. Pronotum yellow with dark brown anterior margin and posterior, inverted U-shaped mark. Episternum bicolor, anteriorly dark brown, and posteriorly yellow; postfurcasternum yellow with dorsal, dark brown area. Mesonotum yellow with dark brown anterior margin; lateral regions with C-shaped, dark brown marks; medial region with a triangular, dark brown mark. Metanotum yellow with lateral, dark brown areas; pteropleura yellow, with dark brown suffusions adjacent to pleural sutures. Foreleg. Coxa yellow with interspersed yellow and black setae; trochanter yellow with brown dorsal surface. Femur posterior surface yellow with small brown areas on base, medial region, and apex; anterior surface yellow with medial, dark brown mark; tibia pale brown with yellow areas at base; basitarsus mostly pale brown, with proximal yellow area on posterior surface; second to fourth tarsomere pale brown. Mid- and hind legs. Mid-leg with yellow coxa, trochanter, and femur; tibia and tarsus pale brown. Hind leg with yellow coxa, trochanter yellow with pale brown; femur yellow with medial, pale brown ring; tibia yellow with posterior ½ and preapical region pale brown; tarsus pale brown. Wings. Forewing membrane pale amber, with slightly marked areas on distal ½ of costal field, subcostal, and radial fields, and on the area surrounding 1m-cu, 2m-cu and 1cua-cup, fork of MP, and stems of RP branches; pterostigma pale brown; venation mostly pale brown with dark areas at the base of R+M, Cu, and anal veins. Hind wing hyaline, with costal and subcostal field pale amber; pterostigma pale brown, venation pale brown, with darker areas near wing base. Abdomen. Tergites pale brown with yellow area adjacent to posterior margins. Sternites yellow, except those of the abdominal segments III-V pale brown.

Figure 22. 

Anchieta sophiae Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal (abdomen removed) B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 22). Head. Diamond-shaped in frontal view, rugose; vertexal region domed above compound eyes; coronal suture noticeable; paraocular area concave. Compound eye hemispherical, as wide as ½ of interocular distance at toruli level. Antenna moniliform, flagellum with 47–53 flagellomeres, discoidal in shape, those of distal ½ wider, except at apex narrower; all flagellomeres with medial ring of short setae. Maxillary palpus with the first two palpomeres as long as wide, third palpomere 3× as long as wide, fourth slightly shorter than third, fifth as long as fourth; labial palpus with first palpomere 1.5× as long as wide, second 5× as long as wide, third slightly shorter than second palpomere; palpimacula narrowly ovoid. Thorax. Pronotum as long as wide, with groove contiguous to lateral and distal margins; in lateral view, posterior margin slightly raised, the rest of the surface straight; entire surface with scattered, thick setae, arising flush the pronotal surface. Postfurcasternum quadrangular with anteroventral narrow process. Mesonotum as wide as long with scattered fine setae on medial region; metanotum ~ 3× as wide as long, glabrous. Pteropleura covered with abundant, thin setae. Foreleg. Coxa slightly shorter than femur, cylindrical, with abundant, fine and long setae; trochanter subtrapezoidal, densely setose, dorsally with a few long, thick setae; anterior surface with blunt process. Femur robust, setose; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes, proximally with a more developed sub-basal process; adjacent row of thickened setae with globular base reduced to distal ½; anteroventral row of processes reduced to sub-basal, primary process, and apex; adjacent row of thickened setae with globular base present nearly over the entire ventral surface. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae; apical region of anterior surface with a patch of clavate setae. Basitarsus with long lanceolate process; basal ½ with clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid- and hind legs. Mid- and hind leg covered with abundant fine setae; hindtibia slightly widened; on both legs basitarsus 2.5× as long as wide, second tarsomere slightly longer than wide; third and fourth as long as wide; fifth tarsomere 1.5× as long as wide. Wings. Forewing oval, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein forked, with 14 or 15 subcostal veinlets; pterostigma rectangular; subcostal space with single, medially located crossvein; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight with 3–6 veins arising from it, three or four from rarp1; M fused basally with R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork, forming a small trapezoidal cell; six or seven gradate crossveins present; Cu deeply forked, CuA apically branched; CuP proximally angled, approaching A1, forked near separation of M and R; A1 simple, A2 forked. Hind wing notably smaller and narrower than forewing; costal space narrow and reduced, with 5–9 subcostal veinlets; C and Sc fused at 1/3 of wing length; Sc vein abruptly curved posteriad at proximal margin of pterostigma, to merge the RA; subcostal space without crossveins; pterostigma elongated, narrow; radial space with single sigmoid crossvein; gradate crossveins absent; two veins arising from each anterior, radial cell. M vein forked at level of R fork; Cu deeply forked, CuA gently bent, distally forked, first branch simple; CuP distally gently bent anteriorly, near posterior wing margin, with two or three branches; A1 simple, A2 short, simple. Abdomen. Medially widened, tergites quadrangular, without keeled processes; sternites rectangular.

Male genitalia (Fig. 23A–E). Tergite IX notably narrower medially than laterally, lateral margin with posterior concavity. Sternites VIII and IX fused, fusion line moderately distinct; sternite IX hourglass-shaped, dorsally forming a canal. Gonocoxite IX sinuous, longer than gonapophyses X, posterior apex pointed, lacking processes. Ectoproct trapezoidal, narrow; lateral apex convex, transparent; posteroventrally set with a patch 16–19 thick, conical setae. Gonocoxites X ventrally canaliculated, hourglass-shaped, curved; posterior ½ with lateral and dorsal processes; base of gonostylus X thickened, triangular with lateral, basal processes; the rest of the structure whip-shaped, short, with apical portion incurved. Gonapophyses X rod-shaped, with dorsally curved tips; both joined by membrane, forming a V-shaped structure. Gonocoxites XI narrow, U-shaped, medial lobe flattened, rounded; lateral arms slightly straight, anterior apex bent ventrally.

Figure 23. 

Anchieta sophiae Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral.

Female genitalia (Fig. 23F–H). Sternite VII noticeably wider laterally than medially, with posterior margin broadly concave. Tergite VIII slightly narrower medially than laterally, enclosing the spiracle of the segment, lateral margin quadrangular. Gonocoxites VIII and medial part of gonapophyses VIII completely fused, forming an enlarged, chamber-shaped structure; lateral part of gonapophyses VIII as a concave, narrow plate, hidden by the tergite IX + ectoproct. Tergite IX + ectoproct triangular. Gonocoxites IX elongated, sinuous, and narrow, as long as the last five abdominal segments together. Gonapophyses IX not discernible. Bursa copulatrix narrow, short, membranous. Spermatheca complex and coiled; proximal section long and thin, forming several coils; medial section thicker than proximal section, forming several coils; distal section wider than medial section, progressively expanded towards apex, sac-like, forming a convolution; fertilization canal duct, long, spiral-shaped, forming several convolutions; fertilization canal elongated, thin, J-shaped, covered with microfilaments.

Distribution

French Guiana (Camopi, Maripasoula).

Remarks

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.

Anchieta tinctus Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 24, 25

Type locality

Costa Rica, Heredia: Est. Biol. La Selva, Arboleda, 10°26'N, 84°01'W, 50–150 m, 28 Jan. 1998, R. Vargas C. leg.

Material examined

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”, MNCR. Paratypes. Costa Rica – Heredia • 1 ♀; Est. Biol. La Selva, Arboleda; 10°26'N, 84°01'W; 50–150m; 28 Jan. 1998; R. Vargas C. leg.; INBIOCRI002286797; MNCR. – Limón • 1 ♂; Sector Cocori, 30 Km al N. de Cariari; 100 m; Jan. 1993; E. Rojas leg.; L-N 286000, 567500, 1753, CRI001675541; MNCR.

• 1 ♂; Sector Cocori; 150 m; Feb. 1993; E. Rojas leg.; L-N 286000, 567500, CRI001309729; MNCR. • 1 ♂; Manzanillo, RNFS Gandoca y Manzanillo; 0–100 m; 20–30 Nov. 1992; F.A. Quesada leg.; L-S 398100, 610600, CRI000784237; MNCR.

Diagnosis

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.

Etymology

The specific epithet of this species is a noun in apposition in allusion to the wing coloration pattern of this species.

Description

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. 24). Head. Mostly dark brown; region of the vertex dark brown with lateral rows of dark brown setae. Postgena pale brown. Antennal scape and pedicel dark brown, flagellum dark brown, except for seven yellow preapical flagellomeres. Frons pale brown. Palpimacula brown. Thorax. Pronotum dark brown with black bristles; episternum and postfurcasternum dark brown. Meso- and metanotum dark brown, mesonotum with dark brown bristles. Pteropleura dark brown. Foreleg. Coxa dark brown with paler apex, covered with dark brown to black setae; trochanter dark brown. Femur mostly dark brown; tibia dark brown, clavate setae pale brown; basitarsus pale brown, with amber apex, second to fourth tarsomere pale brown. Mid- and hind leg. Mid-leg with coxa, trochanter and femur dark brown; tibia mostly dark brown, apex pale brown, entire surface with dark brown to black setae; tarsus pale brown. Hind leg with coxa, trochanter and femur brown, with dark brown setae; outer surface of tibia with base, posterior margin and preapical region dark brown, the rest of the surface including the apex pale brown to orange; inner surface with same pattern as outer surface, sometimes covered with dark brown to black setae; tarsus pale brown. Wings. Forewing membrane mostly pale amber, with darker areas on basal ½ and area adjacent to proximal part of pterostigma; with 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 dark amber; pterostigma brown, becoming paler towards distal region; venation uniformly brown. Hind wing mostly hyaline with area adjacent to posterior and distal margin pale amber, base of wing brown, costal space and base of subcostal field amber; brown pterostigma, uniformly brown venation, dark brown setation. Abdomen. Brown with dark brown margins of sclerites; pleural membrane dark brown.

Figure 24. 

Anchieta tinctus Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal B wings C head, frontal view D pronotum, dorsal E forelegs F hind tibia, outer surface.

Morphology (Fig. 24). Head. Diamond-shaped in frontal view, rugose; vertexal region domed above compound eyes, with lateral rows of short, reclined setae; coronal suture distinct; paraocular area concave. Antenna moniliform, flagellum with 51 flagellomeres, discoidal in shape, those of apex narrower; all flagellomeres with medial ring of short setae. Compound eye hemispheric, as wide as ½ of the interocular distance at toruli level. Thorax. Pronotum slightly longer than wide, with groove contiguous to lateral and distal margins of pronotum; in lateral view, posterior margin slightly elevated, the rest of the surface is straight; entire surface with abundant thick setae arising flush the pronotal surface. Mesonotum slightly wider than long, with abundant thick and long setae; metanotum ~ 3× as wide as long. Pteropleura covered with abundant fine, long setae. Foreleg. Coxa as long as femur, cylindrical, slightly distally widened, with abundant fine, long setae; trochanter subtrapezoidal, densely setose, dorsally with a tuft of long, thick setae, anterior surface with blunt process covered with long setae. Femur robust, with abundant fine, long setae; closing surface with posteroventral row of integumentary specializations fully developed, composed of tubercle-shaped processes with conical Stitz organs (process/seta ratio 1:1 or 2:1), proximally with spine-shaped, sub-basal process (process/seta ratio 4: 1); adjacent row of thickened setae with globular base reduced to distal ¼; anteroventral row of specializations reduced to proximal region and apex, composed of tubercle-shaped processes, with conical setae (process/seta ratio 1:1); basal, primary process present, spine-shaped, curved (process/seta ratio 7:1); adjacent row of thickened setae with globular base present on distal 4/5. Tibia almost as long as femur, curved, glabrous, closing surface with a row of prostrate setae, with patch of clavate setae apically on anterior surface. Basitarsus with long lanceolate process, proximal ½ with clavate setae on anterior surface, ventrally with a row of prostrate setae. Mid- and hind leg. Mid-leg with coxa, trochanter and femur set with long and fine setae, tibia moderately medially expanded, with abundant long and thick setae, tarsus covered with short setae, basitarsus as long as the next three together. Hind leg with coxa, trochanter and femur covered with long, fine setae; tibia conspicuously expanded and laterally flattened, oar-shaped, densely setose; tarsomeres with short, thin setae, basitarsus as long as the next three tarsomeres together. Wings. Forewing oval, venation setose, trichosors present along wing margin except at base; costal space slightly widened medially, humeral vein sometimes forked, with 11–14 subcostal veinlets; pterostigma rectangular; subcostal space with single crossvein, medially located; Sc vein abruptly bent posteriad at proximal margin of pterostigma to merge the RA; rarp2 straight with three or four veins arising from it, three or four from rarp1; M fused basally to R; RP base located near separation of M and R, M fork near such separation; 1r-m located between RP base and M fork forming a small trapezoidal cell; seven or eight gradate crossveins present; Cu deeply forked, CuA apically branched, CuP proximally angled, approaching A1, forked slightly before the level of separation of M and R; A1 simple, A2 forked. Hind wing notably smaller and narrower than forewing, oval; costal space narrow and reduced, with 7–9 crossveins; C and Sc fused at 1/3 of wing length, subcostal space without crossveins; Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge the RA; pterostigma elongated, slightly distally expanded; radial space with single crossvein, sinuous; two or three veins arising from rarp1, one or two from rarp2. M forked at or slightly before R fork. Cubitoanal space with distal crossvein. Cu vein deeply forked, CuA slightly concave, distally forked, first branch simple; CuP distally anteriorly bent near wing margin, with two or three branches; A1 arched A2 short, simple. Abdomen. Medially widened, tergites of abdominal segments III and IV with posteromedial keeled processes. Sternites covered with long, fine setae, gradually becoming longer towards the last abdominal segments.

Male genitalia (Fig. 25A–E). Tergite IX remarkably narrower medially than laterally, lateral region expanded, with blunt process. Sternites VIII and IX fused, with weakly marked fusion line; sternite VIII setose; sternite IX quadrangular with blunt corners, dorsally canaliculate, curved ventrally. Gonocoxites IX reduced, present as fusiform, helical, short and thin sclerites, ventrally attached to lateral arms of gonocoxites XI. Ectoproct trapezoidal, ventrolaterally produced into a rounded, smooth, glabrous outgrowth; ventral surface concave with slightly sclerotized plate on inner side, set with short, thin setae on posterior ½. Gonocoxites X forming a flattened, medially constricted sclerite; anterior ½ expanded, spatulate, posterior ½ triangular, straight; gonostyli X with thickened base, set with short lateral processes articulated to gonocoxites X, medially bent posterodorsally, canaliculated on dorsal surface; the rest of the structure dorsally curved, whip-shaped, forming an apical loop. Gonapophyses X joined by membrane, parallel ventral view; elongated, curved, narrow, with anterior apex spatulate, bent laterally; posterior apex fused to form a long blunt process, dorsally projected. Gonocoxites XI narrow, U-shaped, medial lobe flattened, rounded, medially less sclerotized; lateral arms of gonocoxites XI short, thin, anterior apex bent ventrally.

Figure 25. 

Anchieta tinctus Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca.

Female genitalia (Fig. 25F–H). Sternite VII (gonocoxites VII) as two trapezoidal plates joined by posteromedial bridge; posterior margin broadly concave. Tergite VIII medially narrower than laterally, enclosing spiracle of the segment, lateral margin quadrangular. Gonocoxites VIII as subrectangular, concave, sclerotized plate; gonapophyses VIII medial part composed of quadrangular plates forming a canal, whose ventral surface has longitudinal depression; lateral part of gonapophyses VIII a trapezoidal plate. Tergite IX + ectoproct elongated triangular. Gonocoxites IX elongated, curved and narrow, as long as the last four abdominal segments together. Gonapophyses IX as two small, oval sclerites located behind the gonocoxites IX base. Bursa copulatrix is thin, moderately sclerotized, remarkably long, forming a loop. Spermatheca long, complex and coiled; proximal section narrow, forming three convolutions; medial section gradually expanded, wider than proximal section, forming few convolutions; distal section markedly expanded, wider than medial section, sac-shaped. Fertilization canal duct long, spiral-shaped, forming three convolutions, with preapical widening; fertilization canal elongated, narrow, J-shaped, covered with microfilaments.

Distribution

Costa Rica (Heredia, Limón).

Remarks

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 Navás, 1928

Plega Navas, 1928: 326. Type species: Symphrasis signata Hagen, by original designation.

Further description

Parker and Stange (1965): 605; Penny (1982a, b): 421; Penny and da Costa (1983): 617.

Taxonomy

Parker and Stange (1965): 605 (divided Plega into two groups based on type of antenna and host selection); Werner and Butler (1965): 67 (divided Plega into two groups based on type of antennae which corresponds to host selection); Tjeder (1968): 16–17 (Plega = Trichoscelia).

Key to species

Rehn (1939b): 240–241 (U.S. species); Penny (1982a): 421–422 (Brazilian Amazon); Penny and da Costa (1983): 617 (Brazil).

List of species

Penny (1977): 37; Ardila-Camacho et al. (2019): 352.

Diagnosis

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.

Description

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.

Included species

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)

Biology

Summarized in Ardila-Camacho et al. (2021b).

Etymology

Uncertain, and not specified in the original description of the genus by Navás (1928).

Key to the species of Plega

1 Forefemur remarkably elongated and narrow in lateral view (Fig. 63F) 2
Forefemur narrow and short or moderate- to strongly robust in lateral view (Fig. 32F) 5
2 Forefemur closing surface with anteroventral row of integumentary specializations fully developed (Fig. 63E, F) 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. 77A) Plega zikani Navás, 1936
Antennal flagellum uniformly brown 4
4 Antennal flagellum with basal flagellomeres as wide as long (Fig. 60E) Plega paraensis Penny, 1982
Antennal flagellum with basal flagellomeres 3× as wide as long and subconical (Fig. 38C) Plega duckei Penny, 1982
5 Posterior surface of forefemur pale with numerous brown dots (Fig. 26F) 6
Posterior surface of forefemur pale with brown areas, which may or may not be interconnected (Fig. 28F) 17
6 Pale medial area of forewing pterostigma with anterior notches (Fig. 44B); posteroventral region of male ectoproct with a flattened lobe covered with clear, conical setae (Fig. 45A) 7
Pale medial area of pterostigma with different pattern, or if notched, additional smaller pale areas are present (Fig. 61B); male ectoproct lacking posteroventral flattened lobe with clear, conical setae 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. 45C–E) 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. 31C–E) Plega bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov.
8 Supraantennal region unmodified, without lateral protuberant areas (Fig. 48C, D); male gonocoxite IX posterior apex bifid (49C–E); male ectoproct posterior surface set with abundant, stout, dark setae (Fig. 49A) Plega lachesis Ardila-Camacho & Contreras-Ramos, sp. nov.
Supraantennal region with lateral protuberances covered with reclined setae (Fig. 69C); male gonocoxite lacking digitiform processes or with more than two processes; male ectoproct posterior surface with unmodified setae or with setae arising from protuberant or spiniform bases (Figs 70A, 72A) 9
9 Male gonocoxites IX lacking digitiform processes (Fig. 70C–E) 10
Male gonocoxites IX se with 3–12 digitiform processes 15
10 Antennal flagellum completely brown (Fig. 34A); forefemur anterior surface entirely dark brown (Fig. 34E); male ectoproct lacking modified setal bases (Fig. 35A) 11
Antennal flagellum with 3–5 pale, preapical flagellomeres (Fig. 50D); forefemur anterior surface with base (area adjacent to primary process) pale, brown on the rest (Fig. 50E); male ectoproct bearing modified setal bases on posteroventral surface (Fig. 51A) 12
11 Pale area of pterostigmata enlarged, cream colored (Fig. 56B); forewing generally with a dark brown mark extended from 1m-cu to apical part of intracubital space forming an obtuse angle (Fig. 56B); male sternite IX with thickened, black setae on preapical area (Fig. 57B) Plega obtusa Ardila-Camacho & Contreras-Ramos, sp. nov.
Pale area of pterostigmata small (Fig. 34B); forewing with strong disruptive pattern, with pale wing tips (Fig. 34B); male sternite IX with thin setae over the entire surface (Fig. 35B) Plega disrupta Ardila-Camacho & Contreras-Ramos, sp. nov.
12 Forewing hyaline (Fig. 58B); sternite IX broadly rounded (Fig. 59B); male ectoproct with pedicellate setae on posteroventral area (Fig. 59A) Plega oswaldi Ardila-Camacho & Contreras-Ramos, sp. nov.
Forewing with area adjacent to crossveins and area between apical twigging amber (Fig. 26B); sternite IX trapezoidal (Fig. 27B); male ectoproct with prominent spiniform setal bases on the posteroventral area (Fig. 27A) 13
13 Area adjacent to frontal sutures strongly sunken (Fig. 69C); antennal scape short, lacking stout setae (Fig. 69C, D); male genitalia with ventral process of the median lobe of the gonocoxites XI short and blunt (Fig. 69C–F) Plega spinosa Ardila et al., 2019
Area adjacent to frontal sutures slightly sunken (Fig. 26D); antennal scape elongate set with stout setae in the male (Fig. 26C, D); male genitalia with ventral process of the median lobe of the gonocoxites XI acuminate (Fig. 27F) 14
14 Antennal scape 2× as long as wide at middle (Fig. 26D); hind wing without conspicuous apical stripe (Fig. 26B); male genitalia with ventral part of gonocoxites XI median lobe set with ventral triangular process and caudal transversal ridge (Fig. 27F); female gonocoxites + gonapophyses VIII with narrow and elongate medial process (Fig. 27J) Plega acevedoi Ardila-Camacho & Contreras-Ramos, sp. nov.
Antennal scape 3.5× as long as wide at middle (Fig. 50D); hind wing with conspicuous apical stripe (Fig. 50B); male genitalia with ventral part of gonocoxites XI median lobe set with ventral elongate hook-shaped process and caudal, lateral fins (Fig. 51C–F); female gonocoxites + gonapophyses VIII with vase-shaped medial process (Fig. 51J) Plega longicornis Ardila-Camacho & Contreras-Ramos, sp. nov.
15 Forefemur remarkably robust (Fig. 61F); male gonocoxite IX set with three apical digitiform processes, subequal in shape and size, situated on inner surface (Fig. 62C–E); bursa copulatrix entirely membranous (Fig. 62H Plega pseudohagenella Ardila-Camacho & Contreras-Ramos, sp. nov.
Forefemur not remarkably robust (Fig. 71F); male gonocoxite IX set with 4–12 processes; bursa copulatrix with proximal region sclerotized (Fig. 74I) 16
16 Head with vertexal region patterned with narrow pale and brown stripes (Fig. 71C); pronotum brown with pale medial areas (Fig. 71D); forefemur anterior surface brown with pale areas including the base (Fig. 71E); male gonocoxite IX with 10–12 subequal processes arranged in a bundle (Fig. 72C–E) Plega stangei Ardila et al. 2019
Head with vertexal region with lateral broad brown stripes (Fig. 73C); pronotum mostly brown (Fig. 73D); forefemur anterior surface uniformly brown (Fig. 73E); male gonocoxite IX with 4–6 subequal processes on distal ⅓ arranged as a row of branches on the outer surface (Fig. 74C–E) Plega vangiersbergenae Ardila-Camacho, sp. nov.
17 Basal antennal flagellomeres as wide as long (Fig. 36C) 18
Basal antennal flagellomeres > 2× as wide as long (Fig. 42C) 19
18 Antennal flagellum entirely brown (Fig. 36C, D); supra-antennal area entirely brown (Fig. 36C); male gonocoxite IX with three digitiform, apical processes, whose surface has scattered, minute spinules (Fig. 37C–E); male sternite IX without short, posteromedial process (Fig. 37A); female gonocoxites VII as two lateral trapezoidal plates (Fig. 37G) Plega drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov.
Antennal flagellum with 3–5 pale preapical flagellomeres (Fig. 75C); supraantennal area with a U-shaped pale area (Fig. 75C); male gonocoxite IX with two digitiform, smooth processes, of which one is slightly longer (a third smaller process is sometimes present) (Fig. 76C–E); male sternite IX with short, posteromedial process (Fig. 76A); female gonocoxites VII unpaired, sternite-like (Fig. 76G) 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. 42F) 20
Forecoxa with thickened setae arising from protuberant bases; forefemur not noticeably robust (Fig. 52F) 21
20 Body mostly dark reddish brown (Fig. 42A); forefemur anterior surface uniformly dark-reddish brown colored (Fig. 42E); male gonocoxite IX set with 7–9 long thin processes arranged as a paint-brush (Fig. 43C–E) Plega fumosa Linsley & MacSwain, 1955
Body with a mixture of dark brown and yellow (Fig. 54A); forefemur anterior surface dark brown with base yellow (Fig. 54A); male gonocoxite set with 8–18, short digitiform processes arranged as a paint-brush (Fig. 55C–E) Plega mixteca Ardila et al. 2019
21 Anteroventral row of integumentary specializations lacking curved, basal, primary process (Fig. 46E); postfurcasternum paired, plate-shaped Plega insolita Ardila-Camacho & Contreras-Ramos, sp. nov.
Anteroventral row of integumentary specializations with curved, basal, primary process (Fig. 32E); postfurcasternum unpaired, collar-like 22
22 Basal antennal flagellomeres > 3× as wide as long (i.e., discoidal) (Fig. 28D) 23
Basal antennal flagellomeres 2× as wide as long (Fig. 65C) 24
23 Body color pattern pale and dark reddish brown (Fig. 28A); mesonotum often with a wide, pale, medial band; sternite IX posteromedially elongated, surpassing posterior the margin of ectoproct (Fig. 29A); male gonocoxite IX equipped with 4–7 processes of different lengths arranged as a paint-brush (Fig. 29C–E); female gonapophyses VIII with chamber-shaped medial part, posteromedially set with an elongated Y-shaped process (Fig. 29H) Plega banksi Rehn, 1939
Body color pattern pale and dark brown (Fig. 32A); mesonotum nearly completely dark brown; male sternite IX posteromedially short, not surpassing the posterior margin of ectoproct (Fig. 33A); male gonocoxite IX equipped with 4–15 digitiform processes twisted (Fig. 33C–E); female gonapophyses VII medial part keel-shaped, with short posteromedial, bilobed process (Fig. 33H) Plega dactylota Rehn, 1939
24 Region of the vertex often with a well-defined, arrow-shaped, pale area surrounding the coronal suture (Fig. 52C); pronotum as long as wide and mostly dark brown (Fig. 67D); male sternite IX posteromedially elongated (Fig. 68B) 25
Region of the vertex with pale area surrounding the coronal suture not arrow-shaped (Fig. 65C); pronotum slightly longer than wide and with conspicuous pale areas and stripes (Fig. 65D); male sternite IX posteromedially barely emarginated (Fig. 66B) 26
25 Forewing distinctively enlarged (Fig. 52A); male gonocoxite IX long and thin, equipped with 4–6 processes of different sizes, arranged as a hand (Fig. 53C–E) Plega megaptera Ardila-Camacho & Contreras-Ramos, sp. nov.
Forewing not distinctively enlarged (Fig. 67A); male gonocoxite IX short and thickened, equipped with two or three processes, of which one is apical and longer and 1 or 2 are preapical and shorter (Fig. 68C–E) Plega sonorae Ardila et al., 2019
26 Forefemur anterior surface completely dark brown (Fig. 65E); male gonocoxite IX equipped with 3–5 elongated, intertwined processes (Fig. 66C–E) Plega signata (Hagen, 1877)
Forefemur anterior surface brown with pale base (Fig. 40E); male gonocoxite IX posteriorly, laterally curved, and set with 8–13 processes arranged in a bunch (Fig. 41C–E) Plega flammata Ardila-Camacho & Contreras-Ramos, sp. nov.

Plega acevedoi Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 26, 27

Type locality

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.

Material examined

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; CNIN. Paratypes. Mexico • 1 ♂; Morelos, CEAMISH, Sierra de Huautla; 14 Apr. 1996; S. Zaragoza leg.; CNIN. • 1 ♀; Tlaquiltenango, Huautla; 18°23'04.2"N, 99°03'00.03"W; 1028 m; 25 Nov. 2008; V.H. Toledo; selva baja caducifolia; light trap; CNIN-SHM120. • 1 ♀; Tlaquiltenango, Huaxtla; 18°23'4.272"N, 99°3'0.036"W; 1028 m; 25 Nov. 2008; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN- SHM174. • 1 ♀; Tlaquiltenango, Huaxtla; 18°23'4.272"N, 99°3'0.036"W; 1028 m; 23 Mar. 2009; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN- SHM111.

Other material

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; CNIN-SHM227. • 1 ♂; same data as for preceding; CNIN-SHM229. • 1 ♂; same data as for preceding; CNIN- SHM268. • 1 ♂; same data as for preceding; CNIN-SHM261. • 1 ♂; same data as for preceding; CNIN- SHM262. • 1 ♂; same data as for preceding; CNIN- SHM246. • 1 ♂; same data as for preceding; CNIN- SHM249. 1 ♂; same data as for preceding; CNIN-SHM257. • 1 ♂; same data as for preceding; CNIN- SHM261. • 1 ♂; same data as for preceding; CNIN- SHM253. • 1 ♂; same data as for preceding; CNIN- SHM278. • 1 ♂; same data as for preceding; CNIN- SHM251. • 1 ♂; same data as for preceding; CNIN- SHM195. • 1 ♂; same data as for preceding; CNIN-SHM266. • 1 ♂; same data as for preceding; CNIN-SHM242. • 1 ♂; same data as for preceding; CNIN-SHM193. • 1 ♂; same data as for preceding; CNIN-SHM228. • 1 ♂; same data as for preceding; CNIN-SHM232. • 1 ♂; same data as for preceding; CNIN-SHM241. • 1 ♂; Tlaquiltenango, Huautla, 18°23'04.27"N, 99°03'00.03"W; 1028 m; 25 Nov. 2008; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN-SHM197. • 1 ♂; same data as for preceding; CNIN-SHM208. • 1 ♂; same data as for preceding; CNIN- SHM198. • 1 ♂; same data as for preceding; CNIN-SHM204. • 1 ♂; same data as for preceding; CNIN- SHM265. • 1 ♂; same data as for preceding; CNIN-SHM219. • 1 ♂; same data as for preceding; CNIN-SHM206. • 1 ♂; same data as for preceding; CNIN- SHM272. • 1 ♂; same data as for preceding; CNIN-SHM203. • 1 ♂; same data as for preceding; CNIN-SHM244. • 1 ♂; same data as for preceding CNIN-SHM244. • 1 ♂; same data as for preceding; CNIN-SHM191. • 1 ♂; same data as for preceding; CNIN-SHM269. • 1 ♂; same data as for preceding; CNIN-SHM277. • 1 ♂; same data as for preceding; CNIN-SHM207. • 1 ♂; same data as for preceding; CNIN-SHM209. • 1 ♂; same data as for preceding; CNIN-SHM202. • 1 ♂; same data as for preceding; CNIN-SHM234. • 1 ♂; same data as for preceding; CNIN-SHM210. • 1 ♂; same data as for preceding; CNIN-SHM201. • 1 ♂ same data as for preceding; (CNIN-SHM279). • 1 ♂; same data as for preceding; CNIN-SHM223. • 1 ♂; same data as for preceding; CNIN- SHM199. • 1 ♂; Tlaquiltenango, Huaxtla; 18°23'8.412"N, 99°3'4.068"W; 1023 m; 28–31 Jan. 2009; V.H. Toledo leg.; CNIN-SHM285. • 1 ♂; Tlaquiltenango Huaxtla; 18°23'4.272"N, 99°3'0.036"W; 1028 m; 23 Mar. 2009; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN-SHM167. • 1 ♀; same data as for preceding; CNIN-SHM177. • 1 ♂; Tlaquiltenango, E. Huaxtla; 18°22'50.556"N, 99°2'39.624"W; 1030 m; 29 Mar. 2009; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN. • 1 ♀; Tlaquiltenango, Huaxtla; 18°23'4.272"N, 99°3'0.036"W; 1028 m; 25 Nov. 2008; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN-SHM115. • 1 ♀; same data as for preceding; CNIN- SHM142. • 1 ♀; same data as for preceding; CNIN-SHM144. • 1 ♀; same data as for preceding; CNIN-SHM150. • 1 ♀; same data as for preceding; CNIN-SHM119. • 1 ♀ same data as for preceding; CNIN-SHM152. • 1 ♀; same data as for preceding; CNIN-SHM156. • 1 ♀; same data as for preceding; CNIN-SHM189. • 1 ♀; same data as for preceding; CNIN-SHM166. • 1 ♀; same data as for preceding; CNIN-SHM154. • 1 ♀; same data as for preceding; CNIN- SHM181. • 1 ♀; Tlaquiltenango, Huaxtla; 18°23'8.412"N, 99°3'4.068"W; 1023 m; 28–31 Jan. 2009; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN-SHM134. • 1 ♀; same data as for preceding; CNIN- SHM176. • 1 ♀; Tlaquiltenango, Coaxitlán; 18°27'29.772"N, 99°13'13.872"W; 842 m; 29 Mar. 2011; V.H. Toledo; selva baja caducifolia; light trap; CNIN-SHM185. • 1 ♀; Tlaquiltenango, 2 Km NW Huixastla; 18°28'52.428"N, 99°8'37.032"W; 767 m; 12 Feb. 2010; N. Hernández leg.; selva baja caducifolia; light trap; CNIN-SHM121. • 1 ♀; same data as for preceding; CNIN-SHM127. • 1 ♀; Tlaquiltenango, Huautla; 18°23'04.2"N, 99°03'00.03"W; 1028 m; 25 Nov. 2008; V.H. Toledo leg.; selva baja caducifolia; light trap; CNIN-SHM168. • 1 ♀; same data as for preceding; CNIN-SHM145. • 1 ♀; same data as for preceding; 18°23'08.4"N, 99°03'04"W; 1023 m; 28–31 Jan. 2009; CNIN- SHM147. • 1 ♀; Tepalcingo, S. El Limón; 18°31'42.6"N, 98°56'24.6"W; 1244 m; 09 Feb. 2013; I.A. Villanueva, V.H. Toledo, R. Reyes, J.G. Martínez leg.; selva baja caducifolia; light trap; CNIN-SHM164. • 1 ♀; Puente de Ixtla, Sur de Coaxitlán, camino antiguo a Quetzalapa; 18°25'46.992"N, 99°10'45.408"W; 1050 m; 03 Mar. 2011; V. Toledo, J.G. Martínez, V. Rendón, F. Hinterholzen leg.; selva baja caducifolia; light trap; CNIN-SHM284. • 1 ♀; same data as for preceding; CNIN- SHM109. • 1 ♀; same data as for preceding; CNIN-SHM108. • 1 ♀; same data as for preceding; CNIN-SHM110. • 1 ♀; same data as for preceding; CNIN-SHM122. • 1 ♀; same data as for preceding; CNIN-SHM132. • 1 ♀; same data as for preceding; CNIN-SHM118. • 1 ♀; Tlaquiltenango, Coaxitlán; 18°25'46.992"N, 99°10'45.408"W; 876 m; 03 Mar. 2011; V.H. Toledo, J.G. Martínez, V. Rendón, F. Hinterholsen leg.; selva baja caducifolia; light trap; CNIN-SHM137. • 1 ♀; Tlaquiltenango, 2 Km N Huaxtla; 18°23'27.024"N, 99°2'54.096"W; 1142 m; 13 Nov. 2009; Campos, Reza & Martínez leg.; selva baja caducifolia; light trap; CNIN-SHM126. • 1 ♀; Tlaquiltenango, 5 Km NW de Huaxtla; 18°24'25.524"N, 99°3'14.544"W; 1246 m; 15 Nov. 2009; E.M. Reza, J.G. Martínez, N. Campos, Y. Viveros leg.; selva baja caducifolia; light trap; CNIN-SHM146. • 1 ♂; Tlaquiltenango, 2.5 Km W Huautla, Estación CEAMISH; 18°27'49.28"N, 99°02'15.05"W; 940 m; 15 Feb. 1996; S. Zaragoza, F. Noguera, E. Ramírez & E. Gonzáles leg.; bosque tropical caducifolio; light trap; CNIN. • 1 ♀; Tlaquiltenango, 2.5 Km W Huautla, Estación CEAMISH; 18°27'47.51"N, 99°02'07.90"W; 940 m; 15 Jan. 1996; S. Zaragoza, F. Noguera, E. Ramírez & E. Gonzáles leg.; bosque tropical caducifolio; light trap; CNIN-MDM0123. • 1 ♂; Villa de Ayala, Rancho El Polvorín; 29 Jan. 1971; H. Pérez R. leg.; nocturnal collection; CNIN-N° Cat 86629. • 1 ♂; same data as for preceding; 22 Jul. 1971; CNIN. • 1 ♂; same data as for preceding; 22 Jul. 1971; CNIN. • 1 ♂; same data as for preceding; 17 Oct. 1971; CNIN. • 1 ♂; Villa de Ayala, Polvorín; 02 Oct. 1972; H. Pérez leg.; CNIN. • 1 ♂; Villa de Ayala, Rancho el Polvorín; 1972; H. Pérez R. leg.; nocturnal collection (23:00 h–01:00 h); CNIN. • 1 ♂; same data as for preceding; CNIN. • 1 ♂; Villa de Ayala, Rancho el Polvorín; H. Pérez R. leg.; nocturnal collection (23:00 h–01:00 h); CNIN) • 1 ♀; Villa de Ayala, Rancho El Polvorín, 22 Jul. 1971, H. Pérez leg.; nocturnal collection; CNIN. • 3 ♀; same data as for preceding; 19 Sep. 1971; CNIN. • 2 ♀; same data as for preceding; 1972; 23:00 h–1:00 h; CNIN. • 1 ♀; same data as for preceding; 22 Jul. 1971; CNIN. • 1 ♀; same data as for preceding; 17 Oct. 1971; CNIN. • 2 ♀; Villa de Ayala, Rancho El Polvorín; H. Pérez leg.; nocturnal collection; CNIN. • 1 ♀; same data as for preceding; 20 Jan.1972; nocturnal collection19:00 h–21:00 h; CNIN. • 2 ♀; same data as for preceding; 06 Nov. 1972; CNIN. • 1 ♂; Xochicalco pyramid; 30 Mar. 1962; F.D. Parker & L.A. Stange leg.; FSCA. – Puebla • 1 ♂; Jolalpan, Rancho El Salado, ladera W del cerro Colorado; 18°20'11.61"N, 98°58'55.38"W; 1022 m; 30 Jan. 2014; V.H. Toledo, F. Hinterholzer, J.G. Martínez leg.; selva baja caducifolia; CNIN-SHM226. • 1 ♂; same data as for preceding; CNIN-SHM280. • 1 ♂; same data as for preceding; CNIN-SHM138. • 1 ♂; Jolalpan, Rancho El Salado; 18°20'49.38"N, 98°59'23.02"W; 923 m; 06 Oct. 2010; V.H. Toledo, F. Hinterholzer, J.G. Martínez leg.; selva baja caducifolia; light trap; CNIN- SHM240. • 1 ♂; same data as for preceding; CNIN-SHM238. • 1 ♂; same data as for preceding; CNIN-SHM200. • 1 ♂; same data as for preceding; CNIN-SHM222. • 1 ♂; same data as for preceding; CNIN-SHM180. 1 ♀; Jolalpan, Rancho El Salado, Ladera W de Cerro Colorado; 18°20'11.616"N, 98°58'55.2"W; 1022 m; 05 Oct. 2010; V.H. Toledo, F. Hinterholzer, J.G. Martínez leg.; selva baja caducifolia; light trap; CNIN-SHM170. – Veracruz • 1 ♂; E. Biología Los Tuxtlas; 20 May. 1971; Symphrasis sp., det. Flint, 1978; CNIN-81206.

Diagnosis

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.

Etymology

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.

Description

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. 26). Head. Vertexal region pale, with lateral brown marks extending from occipital ridge to supra-antennal area, embracing pale areas and forming an inverted V-shaped pattern, with pale brown setae; area adjacent to occipital ridge brown; supra-antennal area brown with pale lateral areas, with pale brown setae; occiput and postgena pale with brown stripes. Antennal scape pale with longitudinal brown stripes, apex of dorsal surface with dark brown setae; pedicel brown; flagellum pale brown changing to brown towards apex, with 1–4 pale, preapical flagellomeres, sometimes completely brown. Frons pale with brown with pale medial area. Clypeus and labrum pale with pale brown areas; mandible pale with dark brown corners, apex dark amber; maxilla pale with brown areas, palpus mostly dark brown; labium pale with brown areas, palpus mostly dark brown, palpimacula pale brown. Thorax. Pronotum mostly brown, with pale areas on posterior region; episternum brown; postfurcasternum brown with pale areas. Mesonotum dark brown, sometimes with anterolateral pale areas; metanotum brown with cream anteromedial, small area; pre-episternum brown; pteropleura with brown marks and pale areas, pale brown setae present. Foreleg. Coxa pale to pale brown, with brown mark at apex of anterior surface, with some dark brown setae at basal region; trochanter with same color as coxa, dorsally with some dark brown setae. Femur posterior surface pale with brown dots; anterior surface dark brown, except basal ¼ pale. Tibia dashed with pale and brown. Basitarsus pale at basal ½, changing to amber towards the apex; remaining tarsomeres pale brown. Mid- and hind leg. Coxa and trochanter pale with brown areas; femora and tibiae pale with brown rings; tarsi pale, distal margins of eutarsal tarsomeres with dark brown setae on plantar surface; remaining surface with pale brown setae. Wings. Forewing mostly hyaline; membrane surrounding forks of longitudinal veins, crossveins, first branch of CuA and apex of CuP amber; posterior and apical margins with intermittent, amber areas between apical branches of longitudinal veins; pterostigma brown with small, anterior pale areas, and larger cream medial area; major veins, subcostal veinlets, and wing margin alternating pale and brown; crossveins dark brown. Hind wing hyaline, adjacent area to posterior and apical margins with intermittent amber areas between apical branches of longitudinal veins; pterostigma brown with small pale areas, and wide, cream, preapical region; longitudinal veins alternating pale and brown; crossveins brown, except sigmoid 1r-m bicolor; wing margin alternating pale and brown. Abdomen. Tergites brown. Pleural membrane dark brown, except near dorsal and ventral margins pale. Sternites pale with lateral, brown stripe on each side.

Figure 26. 

Plega acevedoi Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 26). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct, area adjacent to frontal sutures sunken; ​​supra-antennal raised laterally, with abundant, fine, reclined setae; paraocular area concave. Antenna filiform, short; scape two times as long as wide at middle, dorsal surface with stout setae near distal margin; pedicel slightly longer than wide; flagellum not dorsoventrally flattened, with 28–33 flagellomeres, those of flagellar base as wide as long, then changing to slightly longer than wide on the rest of flagellum; all articles with medial ring of fine, short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus narrow. Labrum pentagonal with thin, short setae; maxillary palpus with first palpomere as long as wide, second 1.2× as long as wide, third palpomere 3.5× as long as wide, fourth palpomere 2.5× as long as wide, fifth palpomere slightly longer than third; mentum with long, thin setae; labial palpus with first palpomere 2× as long as wide, second palpomere four 4.5× as long as wide, third slightly shorter than second, palpimacula narrowly ovoid. Thorax. Pronotum slightly longer than wide, with raised anterior margin, medial and posterior regions; raised regions and anterior and margins covered with pedicellate, thickened setae. Mesonotum 2× as wide as long, with scattered, thick, pedicellate setae on medial area. Metanotum ~ 2× as long as wide, glabrous. Pteropleura with short, thin, setae a few thickened on mesanepisternum. Foreleg. Coxa nearly as long as femur, cylindrical, posterior surface with pedicellate, thick setae; trochanter trapezoidal, with thin and short setae, except on dorsal and anterior surfaces with some thickened, pedicellate setae near apex; anterior surface with protuberant area. Femur robust, covered with abundant, fine, short setae; closing surface with posteroventral row of processes composed two medially located, primary processes, with or without a tertiary process between them; proximal portion of the row with sub-basal secondary process and basal tertiary process; the rest of the row with numerous tubercle-shaped processes, and stinger-shaped setae; distal ½ slightly carinated, composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½; it is composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of a few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ½. Tibia almost as long as femur, curved, with thin, short setae; ventral surface keeled with prostrate setae; a patch of clavate setae apically on anterior surface is present. Basitarsus with lanceolate process reaching the middle of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 7× as long as wide; third tarsomere as long as wide, fourth tarsomere two times as long as wide. Mid- and hind leg. Coxa with short and thin setae; trochanter with short, thin setae; femora with thin setae; tibial spurs short; tibiae with short and thin setae. Hind leg longer than midleg; tibia, 1.5× as long as femur; tarsi with fine and short setae, except on distal margin of plantar surface with lateral groups of 3–5 thickened setae; on both legs, basitarsus 4× as long as wide, second tarsomere 1.2× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing oval, trichosors present along margin except on wing base; venation setose; costal space proximally expanded, humeral vein branched, 12–19 subcostal veinlets; pterostigma trapezoidal, elongated, narrow, straight, with distinct veinlets; subcostal space with single crossvein, medially located; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge the RA; radial space with two crossveins; rarp2 gently curved with 2–4 RP branches; 2–4 veins arising from rarp1; M vein basally fused to RA; RP base widely separated from divergence of M and R; M forked opposite to RP origin, 1 r-m connecting RP base and MA stem, forming a trapezoidal cell; 5–7 gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked opposite to the level of separation of M and R; A1 apically forked, ending on posterior margin slightly beyond the level of CuP fork, A2 forked slightly beyond to CuP angle level. Hind wing smaller and narrower than forewing, oval; costal space narrow and reduced, with 5–7 veinlets; C and Sc fused at proximal ⅕ of wing length, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge RA; pterostigma elongated, narrow, straight, composed of well-defined veinlets; radial space with single crossvein, oblique; 2–4 veins arising from rarp1, two or three from rarp2; 1r-m sigmoid, connecting the stems of M and RP. Media forked slightly beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent; CuP not touching A1, strongly anteriorly bent at distal 1/3, pectinate; two crossveins on cubitoanal space present; A1 simple, ending at wing margin at 1r-m stem level; A2 simple, short, and curved. Abdomen. Tergites quadrangular with anterolateral elongated scars; sternites rectangular.

Male genitalia (Fig. 27A–F). Tergite IX medially narrower than laterally; lateral margin blunt. Sternum VIII rectangular; sternite IX trapezoidal in ventral view; posterior margin with medial lobe not developed, but corresponding area dorsally canaliculated; in lateral view trapezoidal, not reaching posterior margin of ectoproct. Gonocoxites IX thin, sinusoid, short; base spatulated; apex pointed, without processes. Ectoproct ovoid, posteroventral surface set with spine-shaped setal bases, remaining surface with long, pedicellate setae; anteroventrally with enlarged, rounded, sclerotized lobe, continuous with ventromedial sclerotized, curved sulcus. Gonocoxites X forming a short, straight, ventrally canaliculate sclerite; anterior apex, expanded; posterior apex with dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X with a membrane; gonostyli X with thickened and concave base with two lateral processes, the rest of the structure, ventrally strongly curved, and posteriorly recurved at mid length before protruding from abdomen, apex abruptly narrowed and pointed. Gonapophyses X rod-shaped, gently curved; gonapophyses sub-parallel, joined by a membrane covering the gonostyli X base. Gonocoxites XI U-shaped, medial lobe complex and elaborated with two differentiated parts: dorsal part as an arch, with lateral parts expanded; ventral part with prominent, rounded, transversal ridge; ventrally a pointed, curved process is present; between these parts an irregular, less sclerotized, hyaline area is present, which forms a triangular outgrowth; lateral arms of gonocoxites XI short, thin, sigmoid, with anterior apex bent ventrad. Hypandrium internum triangular, with lateral fins.

Figure 27. 

Plega acevedoi Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F same, caudal G female terminalia, lateral H same, ventral I spermatheca J gonapophyses VIII of female, ventral.

Female genitalia (Fig. 27G–J). Sternum VII rectangular with posteromedian concavity. Tergite VIII narrower medially than laterally, encircling the spiracle of the segment; lateral margin quadrangular. Gonocoxites VIII not discernible, forming a complex with the gonocoxites VIII. Gonocoxites + Gonapophyses VIII as two lateral quadrangular plates which form a tubular, medial process whose apex is shallowly bifid; lateral part of gonapophyses VIII as a rounded plate folded beneath tergite IX + ectoproct. Tergite IX + ectoproct triangular. Gonocoxites IX elongated and narrow, nearly as long as the last five abdominal segments; gonapophyses IX as tiny sclerites at the inner base of gonocoxites IX. Bursa copulatrix short, funnel-shaped, with proximal-most part lightly sclerotized, the rest membranous and striated. Spermatheca short and simple; proximal and medial sections undifferentiated, thin; proximal-most forming a blunt diverticulum; distal section progressively slightly expanded, forming a blunt diverticulum; fertilization canal duct with proximal portion triangular, concave; fertilization canal elongated a narrow, C-shaped, covered with microfilaments.

Distribution

Mexico (Morelos, Puebla, Veracruz).

Remarks

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

Figs 28, 29

Plega banksi Rehn, 1939: 248. Holotype: male, USA, Arizona (AMNH).

Material examined

Paratypes. USA • 1 ♂; Arizona, Oracle 14 m E.; 27 Jul. 1924; E.P.V. Van Duzee leg.; “Plega banksi Rehn, Paratype”; CAS. • 1 ♀; Baboquivari Mts.; 15 Aug. 1922; O.C. Poling leg.; “Pres. By C.L. Fox”; “Plega banksi Rehn, Paratype”; CAS. • 1 ♂; Boya Thomson, Aboretum, Superior; 1–3 Aug. 1937; MCZ-Paratype 25378.

Other material

Mexico – Sinaloa • 2 ♀; 16 mi S Guamuchil; 20 May. 1962; F.D. Parker & L.A. Stange leg.; Plega fratercula Rehn det. L. Stange, 1991; FSCA. • 1 ♀; Sinaloa, Guamuchil; 01 Jul. 1963; F.D. Parker & L.A. Stange leg.; FSCA. • 1 ♂ 1 ♀; Sinaloa, 16 miles SO. Guamuchil; 16 Jun. 1961; FSCA. • 1 ♂; Sinaloa, Rio Piaxtla, 1 mi. W. Mex No. 15; E.L. Sleeper & R.C. Anderson leg.; Plega dactylota, det. Penny, 1998; CAS. • 1 ♂; Sinaloa, 16 miles SO. Guamuchil; 16 Jun. 1961; F.D. Parker & L.A. Stange leg.; FSCA. – Sonora • 1 ♀; 40 mi N. Guaymas; 29 Aug. 1959; L.A. Stange & A.S. Menke leg.; FSCA. • 1 ♀; 100 yds ½ mi E of San Carlos; 05 Apr. 2003; on Bebbia, REW; UCD. • 1 ♂; Punta La Manga; 27°58'46"N, 111°07'58"W; 5 m; 1–2 Aug. 2009; E.M. Fisher leg.; MT; CSCA.

USA – Arizona • 1 ♂; Madera Canyon; 24 Jul. 1976; B.K. Dozier leg.; FSCA. • 1 ♂; Sta. Catalina Mts.; 01 Jul. 1938; Bryant leg.; FSCA. • 1 ♂; Santa Cruz Co., Peña Blanca, Pajarito Mts.; 07 Jul. 1962; R.H. Arnett Jr., E.R. VanTassell leg.; FSCA. • 1 ♂; Santa Cruz Co., Madera Cyn.; 4880 ft.; 21 Aug. 1963; V.L. Vesterby leg.; UCD. • 1 ♂; same data as for preceding; 15 Aug. 1963; UCD. • 2 ♂; same data as for preceding; 14 Aug. 1963; UCD. • 2 ♂; same data as for preceding; 23 Jul. 1963; UCD. • 2 ♂; same data as for preceding; 15 Aug. 1963; UCD. • 1 ♂; same data as for preceding; 01 Sep. 1963; UCD. • 2 ♂; same data as for preceding; 21 Jul. 1963; UCD. • 1 ♂; same data as for preceding; 12 Sep. 1963; UCD. • 1 ♂; same data as for preceding; 21 Sep. 1963; UCD. • 1 ♂; same data as for preceding; 12 Aug. 1963; UCD. • 7 ♂; same data as for preceding; 21 Jul. 1963; UCD. • 1 ♂; same data as for preceding; 23 Jul. 1963; UCD. • 6♂ 6♀; Tucson, Pima Co.; 02 Aug. 1991; R. & J. Robertson leg.; CAS. • 1 ♂; Pima Co., Bavoquivari Mts., Sabino Can.; 07 Aug. 1951; L.M. Martin leg.; “Phillip Adams Collection 1998 request to Calif. Acad. Sci.”, Plega banksi Rehn”; CAS. • 1 ♀; Globe; 17 Jul. 1933; Parker leg.; Plega fratercula Rehn, det. J.W.H. Rehn, A.N.S.P., 1939; CAS-114. • 1 ♀; Pima Co., Alamo Canyon, Ajo Mts.; Jul. 1924; H.B. Leech & J.W. Green leg.; Plega banksi, det. N. Penny, 1985; CAS. • 1 ♂; Pima Co., Sta. Rita Mts., mouth Madera Canyon Procter Ranch Road; 4400’; 15 May. 1966; R.G. Beard leg.; U.V. light; “Phillip Adams Collection 1998 request to Calif. Acad. Sci.”, Plega banksi Rehn, 1939, det. R.G. Beard; CAS. 1 ♂; same data as for preceding; “LDL#0562”; CAS. • 1 ♀; Pima Co., Sta. Rita Mts., mouth Madera Canyon Procter Ranch Road; 17 Jul. 1966; 4400’; R.G. Beard; U.V. light; MCZ-ENT00681784. • 1 ♂; same data as for preceding; 15 Jul. 1966; Plega banksi Rehn, 1939, det. R.G. Beard; MCZ-ENT00681783. • 14 ♂ 12 ♀; Pima Co., Santa Rita Mts Experimental Range, Florida Cyn.; 31°45.42'N, 110°50.43'W; 1646 m; 24 Apr.–03 Jul. 2014; M.I. Erwin, C. O’ Brien, M.J. Sharkey leg.; Malaise on hillside trail; CSCA. • 1 ♂ 2 ♀; same data as for preceding; 3–10 Jul. 2014; CSCA. • 1 ♂; Pima Co., Sabino Cyn.; 04 Sep. 1961; J.S. Buckett leg., UCD. • 1 ♂; same data as for preceding; 04 Sep. 1961; UCD. • 1 ♂; same data as for preceding; 01 Sep. 1963; V.L. Vesterby; UCD. • 3 ♂; same data as for preceding; 19 Aug. 1963; UCD. • 1 ♂; same data as for preceding; 5800’; 11 Jul. 1967; R.C. Gardner, C.R. Kovacic, K. Lorenzen leg.; UCD. • 2 ♂ 1 ♀; Pima Co., Vail, Mountain Creek Ranch; 32°04.99'N, 110°39.56'W; 1100 m; 06–24 Aug. 2014; M.E. Irwin, M.J. Sharkey leg.; Malaise in small dry wash; CSCA. • 1 ♂; Maricopa Co., Sunflower, Jct. Hwy, 87 & Sycamore Ck.; 33°52'8.616"N, 111°27'51.732"W; ± 100 m; 744 m; 15 Aug. 2012; Oswald, Diehl, Machado leg.; MV light; TAMUIC-TAMU-ENTO X0966102. • 1 ♂; Graham Co., Mt Graham, SE side, 3.5 Km W Jet. 191 on hwy 266; 32°34.32'N, 109°43.11'W; 1265 m; 08–15 May. 2015; Malaise in sandy wash; CSCA. • 1 ♂ 1 ♀; Santa Cruz Co., 10 mi NW Nogales; 04 Sep. 1968; T.R. Haig leg.; ex-fluorescent black light; CSCA. • 1 ♂; Santa Cruz Co., Kino Springs; 20 May. 2002; K.A. & E.E. Williams; CSCA.

Diagnosis

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.

Description

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. 28). Head. Vertexal mostly dark reddish brown with, small, lateral pale areas; area adjacent to occipital ridge pale; area adjacent to coronal suture with paler, arrow-shaped mark; supra-antennal area dark reddish brown; occiput pale with dark brown mark, postgena pale with brown area. Antennal scape brown with pale, longitudinal band on outer surface, pedicel dark brown; flagellum brown. Frons dark reddish brown, sometimes with medial pale stripe. Clypeus pale with lateral brown areas. Labrum pale brown; mandible yellowish with dark brown corners, dark amber at apex; maxilla pale with brown areas, palpus brown, with apex of the last three palpomeres yellowish; labium pale with dark brown postmentum, palpus with first two palpomeres dark brown, the last pale brown; palpimacula pale brown. Thorax. Pronotum pale with medial band and posterolateral dark reddish brown areas; episternum dark reddish brown with yellowish, medial area; postfurcasternum pale. Mesonotum, either mostly dark reddish brown with area adjacent to sutures pale brown or dark reddish brown with wide, pale, medial band with a rhomboid dark mark on center. Metanotum dark brown, sometimes with pale anteromedian area; mesopre-episternum dark reddish brown; pteropleura dark reddish brown with pale areas. Foreleg. Coxa yellowish with reddish brown areas, with interspersed pale and dark brown setae with brown bases; trochanter pale to yellowish with ventral dark reddish brown mark, dorsally with some dark brown setae. Femur posterior surface pale, with widened dark reddish brown areas; anterior surface dark reddish brown, except small, basal, yellowish areas. Tibia posterior surface alternating dark and pale areas, anterior surface mostly dark reddish brown, with preapical yellowish area. Basitarsus with base and lanceolate process dark amber, plus a sub-basal yellowish area; remaining tarsomeres yellowish to pale brown. Mid- and hind leg. Coxae pale with reddish brown suffusions; trochanter pale with brown suffusions. Femora pale with brown sub-basal, ventral area and apex. Mid tibia dashed with pale and brown, hind tibia pale with brown suffusions, the apex brown. Tarsi pale brown. Wings. Forewing mostly hyaline; membrane surrounding crossveins, R fork, RP branches stem, first branch of CuA and apex of CuP, and terminal forks of longitudinal veins amber; a zigzagged amber mark connecting MA and MP forks, terminal part of MA and 2r-m and 1ma-mp, plus an amber area between CuP and posterior wing margin are frequent. Pterostigma brown with pale medial area. Major veins, subcostal veinlets and wing margin alternating pale and dark reddish brown; crossveins dark reddish brown. Hind wing hyaline, amber on area adjacent to stem of first branch of CuA; pterostigma brown with pale, preapical area; longitudinal veins alternating pale and dark brown areas; subcostal veinlets nearly transparent; crossveins brown, 1r-m often bicolor; wing margin alternating pale and brown. Abdomen. Tergites dark reddish brown. Pleural membrane pale with dark reddish brown suffusions. Sternites pale with wide, dark reddish brown, lateral areas, setation mostly pale.

Figure 28. 

Plega banksi Rehn, 1939 A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 28). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct; medial area of ​​supra-antennal region not raised, with fine, reclined setae; paraocular area concave. Antenna sub-moniliform, nearly as long as head and thorax together; scape as long as wide, cup-shaped, ventrally with a few thickened setae; pedicel as long as wide; flagellum not dorsoventrally flattened, with 45–64 flagellomeres, those of proximal ¼ discoidal, then changing from two times as long as wide to as long as wide; all articles with medial ring of fine, short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus narrow. Labrum pentagonal; maxillary palpus with first palpomere as long as wide, second 1.2× as long as wide, third palpomere 4× as long as wide, fourth palpomere 2.5× as long as wide, fifth palpomere slightly longer than third; mentum with long, thin setae; labial palpus with first palpomere two times as long as wide, second palpomere 4× as long as wide, third palpomere slightly longer than second, palpimacula ovoid. Thorax. Pronotum as long as wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae. Mesonotum slightly wider than long, with abundant, thick, pedicellate setae on medial area. Metanotum ~ 2× as wide as long. Pteropleura with short, thin setae, thicker on mesanepisternum and metakatepisternum. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with pedicellate, fine or thick setae of different sizes; trochanter trapezoidal, dorsal surface with some thickened, pedicellate setae near distal margin; anterior surface without protuberant area. Femur robust, closing surface with posteroventral row of processes composed two medially located, primary processes; proximal portion of the row with sub-basal secondary process and basal tertiary process; the rest of the row with abundant tubercle-shaped processes, and stinger-shaped setae; distal portion carinate, composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½; it is composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ⅘. Tibia almost as long as femur, curved, ventral surface keeled with row of prostrate setae; patch of clavate setae apically on anterior surface present. Basitarsus with lanceolate process reaching base of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 9× as long as wide; third tarsomere 1.5× as long as wide; fourth tarsomere two times as long as wide. Mid- and hind leg. Coxa and trochanter with short, thin setae; femora with interspersed fine setae of different lengths and few thickened setae; tibiae mostly with short fine setae; hind leg longer than midleg, tibia 1.5× as long as femur; tarsomeres with distal margin of plantar surface with lateral groups of 5–8 thickened setae; on both legs, basitarsus 3.5 times as long as wide, second tarsomere 1.2× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing narrowly oval, trichosors present along margin except on wing base; costal space proximally expanded, humeral vein branched; 12–15 subcostal veinlets; pterostigma elongated, narrow, straight, with incomplete veinlets; subcostal space with single, medially located crossvein; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge with RA; radial space with two or three crossveins; rarp2 gently curved with three or four RP branches; three or four veins arising from rarp1; M vein basally fused with RA; RP base widely separated from divergence of M and R; M forked opposite to RP origin; 1 r-m connecting RP base and M fork, forming a trapezoidal cell; 3–6 gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked slightly beyond the level of separation of M and R; A1 apically forked, ending on posterior margin at level of CuP fork, A2 forked slightly beyond CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with 5–9 veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge RA; pterostigma elongated, narrow, gently curved, composed of incomplete veinlets; radial space with single crossvein, oblique; three or four veins arising from rarp1, one or two from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent or present; CuP not touching A1, strongly anteriorly bent at distal ½, pectinate, sometimes fused to first branch of CuA; two crossveins on cubitoanal space; A1 apically simple or forked, ending on wing margin at 1r-m stem level; A2 simple, short, and curved. Abdomen. Cylindrical to medially expanded, with subquadrate tergites; tergites III and IV with posteromedian concavity; tergites III-VII with elongated anterolateral scars; sternites rectangular.

Male genitalia (Fig. 29A–E). Tergite IX medially narrower than laterally; lateral margin rounded. Sternite VIII rectangular; sternite IX pentagonal in ventral view, with abundant setae; posterior margin with medial, elongated, blunt lobe which is dorsally canaliculated; in lateral view triangular, apex surpassing posterior margin of ectoproct. Gonocoxites IX short, thickened, sinusoid; base flattened, spatulated; apex laterally curved, branched, with 4–7 processes of different lengths. Ectoproct ovoid, covered with abundant, thickened setae on posterior surface, posteroventrally with blunt lobe; anteroventrally with flattened, rounded lobe that is continuous with ventromedial sclerotized, curved sulcus. Gonocoxites X forming a short, ventrally canaliculated sclerite, whose posterior apex has dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X; gonostyli X with thickened and curved base, equipped with two lateral processes, the rest of the structure, ventrally curved, and anteriorly coiled, forming two loops before protruding from abdomen. Gonapophyses X long, straight, narrow, with posterior apex slightly expanded; gonapophyses arranged in a V-shaped structure, joined by a membrane covering the gonostyli X base. Gonocoxites XI thin, U-shaped, medial lobe complex and elaborated, with two differentiated parts: dorsal part as rounded, anteriorly curved lobe; ventral part with a convex area covered with microspinules, continuous with ventral, curved, prominent hook-shaped process, whose apex is covered with microspinules; between these parts a narrow, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI gently sigmoid, with anterior apex straightly angled. Hypandrium internum triangular.

Figure 29. 

Plega banksi Rehn, 1939 A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 29F–I). Sternite VII rectangular with broad posteromedian concavity connected with medial, narrow, glabrous, sclerotized area; with long, thickened setae near posteromedian concavity. Tergite VIII conspicuously narrower medially than laterally, encircling the spiracle of the segment; lateral margin D-shaped. Gonocoxites VIII as a sub-rectangular, concave sclerite, posteromedially connected to gonapophyses VIII. Gonapophyses VIII with medial part chamber-shaped, ventral surface with lateral convex areas, and lateral surfaces concave, posteromedially with an elongated, Y-shaped process; lateral part of gonapophyses VIII oval. Tergite IX + ectoproct D-shaped. Gonocoxites IX remarkably elongated, sinuous, and narrow, nearly as long as the whole abdomen. Bursa copulatrix short with proximal part strongly expanded, ovoid, and sclerotized, continuous with a membranous and striated part, which is abruptly narrowed. Spermatheca long, complex and entangled; proximal section thin, long, forming numerous coils; medial section as wide as proximal section, forming a long spiral; distal section entangled, as wide as medial section, terminating in an expanded portion, where a blunt diverticulum is present; fertilization canal duct proximally triangular, concave; fertilization canal elongated, pod-shaped, covered with microfilaments.

Distribution

Mexico (Sinaloa, Sonora), USA (Arizona).

Remarks

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 Oswald et al. (2002) and Reynoso-Velasco and Contreras-Ramos (2008) are more likely erroneous and probably represent specimens of the closely related species P. megaptera.

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.

Plega bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 30, 31

Type locality

Ecuador, [Guayas]: Guayaquil, Rosemberg leg.

Material examined

Holotype male, pinned, with genitalia in a separate vial. Original label: “Ecuador, Guayaquil, Rosemberg coll., Acq. 1903”; FSCA.

Diagnosis

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.

Etymology

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.

Description

Measurements. Male (n = 1). Forewing length: 10.4 mm; Hind wing length: 8.3 mm.

Coloration (Fig. 30). Head. Vertexal region pale, with lateral dark brown markings extending from occiput to toruli; supraantennal area dark brown with lateral, small pale areas; pale brown setae present; occiput pale with sark brown spot, postgena pale with brown area. Antennal scape pale brown with dorsal, darker, longitudinal band, entire surface with brown setae; pedicel brown; flagellum brown with three pale, preapical flagellomeres. Frons with dark brown trapezoidal areas below toruli, medial region with pale triangular area. Clypeus pale. Labrum pale brown; mandible pale brown with dark brown corners, amber at apex; maxilla brown with darker areas, palpus pale brown with darker rings on third and fifth palpomere; labium pale brown with darker postmentum, ligula and first two palpomeres; palpimacula pale brown. Thorax. Pronotum medially pale with brown areas, area adjacent to lateral margins dark brown, setae brown, with some paler; episternum pale brown; postfurcasternum pale with posterior area brown. Mesonotum pale with brown areas on the medium and laterally dark brown; metanotum mostly dark brown; pre-episternum brown; pteropleura brown with pale on margins of sclerites, setation mostly pale brown. Foreleg. Coxa pale, with a few, small, brown marks, setae mostly brown, with some paler; trochanter pale with small, brown marks, setation mostly pale brown, with some darker on dorsal and anterior surfaces. Femur posterior surface pale with brown dots, apex brown; anterior surface pale with extensive brown area; processes pale brown. Tibia dashed with pale and dark brown. Basitarsus with base and lanceolate process pale brown, sub-basal area pale, clavate setae pale brown; remaining tarsomeres pale brown. Mid- and hind leg. Mid- and hind coxae mostly brown, setae pale brown; trochanter pale with brown suffusions. Femur of mid-leg and tibia of both legs dashed with pale and dark brown bands, femur of hind leg pale; tibial spurs pale brown. Tarsi pale brown; first four tarsomeres with brown setae, laterally on distal margin of plantar surface; pretarsal claws pale brown. Wings. Forewing mostly hyaline; membrane surrounding crossveins, M fork, first branch of CuA and apex of CuP amber; intermittent amber areas between terminal forks of longitudinal veins on distal ½ of wing; pterostigma brown with wide, pale, medial area; major veins, subcostal veinlets and wing margin alternating pale and brown. Hind wing hyaline; pterostigma brown with wide, pale, preapical area; longitudinal veins alternating pale and brown, subcostal veinlets pale brown; crossveins brown, except sigmoid 1r-m, bicolor; wing margin alternating pale and brown. Abdomen. Cleared.

Figure 30. 

Plega bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal (abdomen removed) B wings C head, frontal D pronotum, dorsal E forefemur, anterior Surface F same, posterior surface.

Morphology (Fig. 30). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct; medial area of ​​supra-antennal region slightly raised, with fine, reclined setae; paraocular area concave. Antenna sub-moniliform, short; scape 1.5× as wide as long, slightly distally expanded, with fine, short setae; pedicel as long as wide; flagellum not dorsoventrally flattened, with 38 flagellomeres, those of proximal ¼ of the flagellum as long as wide; slightly longer than wide on the rest of flagellum; all articles with medial ring of fine, short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus narrow, with fine and short setae. Labrum pentagonal with thin, short setae; maxillary palpus with first palpomere as long as wide, second 1.2× as long as wide, third palpomere 3.5× as long as wide, fourth palpomere 2.5× as long as wide, fifth as long as third, all with short and thin setae; mentum with long, thin setae; labial palpus with first palpomere 2× as long as wide, second palpomere 4× as long as wide, third palpomere slightly longer than second, palpimacula narrowly ovoid. Thorax. Pronotum slightly longer than wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae; remaining surface with fine, short setae. Mesonotum slightly wider than long, with scattered, thick, pedicellate setae on medial area. Metanotum ~ 2× as wide as long, with a few fine, short setae. Pteropleura with short, thin setae. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with abundant fine setae of different lengths; trochanter trapezoidal, with thin and short setae, except on dorsal and anterior surfaces with some thickened, pedicellate setae; anterior surface with protuberant area. Femur robust, covered with abundant, fine, short setae; closing surface with posteroventral row of processes composed two medially located, primary processes, with single secondary process between them; proximal portion of the row with two secondary processes; the rest of the row with abundant tubercle-shaped processes, stinger-shaped setae; distal portion raised composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½; it is composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of a few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ½. Tibia almost as long as femur, curved, with thin, short setae; ventral surface keeled with prostrate setae; a patch of clavate setae apically on anterior surface is present. Basitarsus with lanceolate process reaching the middle of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 7× as long as wide; third tarsomere as long as wide, fourth tarsomere 2.5× as long as wide. Mid- and hind leg. Coxa and trochanter with thin setae, shorter on trochanter; femora and tibiae with fine setae of different lengths; tibial spurs short; hind leg longer than midleg, tibia, 1.5× as long as femur; tarsi with fine and short setae, except on distal margin of plantar surface with lateral groups of three or four thickened setae; on both legs, basitarsus 4× as long as wide, second tarsomere 1.2× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing oval, trichosors present along margin except on wing base; venation setose; costal space proximally expanded, humeral vein branched, 15 or 16 subcostal veinlets, where one or two are forked; pterostigma trapezoidal, straight, with incomplete veinlets; subcostal space with single crossvein, nearly opposite to R fork; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 curved with three RP branches; three veins arising from rarp1; M vein basally fused with RA; RP base not widely separated from divergence of M and R; M forked slightly before RP origin, 1 r-m connecting RP base and M fork, forming a trapezoidal cell; five or six gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked slightly beyond the level of separation of M and R; A1 simple, ending on posterior margin slightly beyond the level of CuP fork, A2 forked slightly beyond CuP angle level. Hind wing smaller and narrower than forewing, oval; costal space narrow and reduced, with 6 veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge RA; pterostigma elongated, slightly subapically expanded, straight, composed of incomplete veinlets; radial space with single crossvein, oblique; three or four veins arising from rarp1, one from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA gently sinuous, first branch candelabrum-shaped, spur vein absent; CuP not touching A1, strongly anteriorly bent at distal 1/3, pectinate; two crossveins on cubitoanal space; A1 simple, ending on wing margin at 1r-m stem level; A2 simple, short, and curved. Abdomen. Cylindrical to medially expanded, setae on tergites, scattered, thin, and short, gradually longer towards terminal segments; tergites subquadrate. Sternites rectangular, with abundant, scattered, thin setae.

Male genitalia (Fig. 31). Tergite IX medially narrower than laterally; lateral margin rounded. Sternite VIII quadrangular; sternite IX pentagonal in ventral view, with rounded posterolateral lobes covered with abundant, long setae; posterior margin with medial, short, triangular lobe which is dorsally canaliculated; in lateral view bluntly trapezoidal, apex not reaching posterior margin of ectoproct. Gonocoxites IX long, gently sigmoid; base flattened, dorsally curved, connected to gonocoxites XI with a membrane; apex straight, thickened, with elongate preapical process on inner surface and seven or eight short processes on outer surface, arranged in a single plane. Ectoproct ovoid, covered with thickened setae on posterior surface, posteroventrally with rounded lobe covered with fine, pedicellate setae; Gonocoxites X forming a short, thickened, ventrally canaliculated sclerite, with anterior apex dorsally bent, posterior apex with dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X with a membrane; gonostyli X with thickened, obtuse base with two lateral processes, the rest of the structure, whip-shaped, ventrally curved, and anteriorly coiled, forming a single loop before protruding from abdomen. Gonapophyses X rod-shaped, straight, posterior apex slightly expanded; gonapophyses arranged in a V-shaped structure joined by a membrane covering the gonostyli X base. Gonocoxites XI thin, U-shaped, medial lobe complex and elaborated, with two differentiated parts: dorsal part as a narrow arch; ventral part forming covering with two lateral concavities, ventrally it is produced as a straight, trapezoidal process; between these parts a narrow, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI gently curved, with anterior apex straightly angled, quadrangular.

Figure 31. 

Plega bowlesi Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral.

Distribution

Ecuador (Guayas).

Remarks

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, 1939

Figs 32, 33

Plega dactylota Rehn, 1982: 423. Holotype: male, USA, Arizona (ANSP).

Plega dactylota lipanica Rehn, 1982: 423. Holotype: male, USA, Texas (ANSP). Rice (1987): 342.

Plega fratercula Rehn, 1939: 247. Holotype: male, USA, Arizona (MCZ), specimen examined. New synonym.

Material examined

Holotype of Plega fratercula

USA • ♂; Arizona, Capitain Mt.; 8 Aug. 1933; R. Anderson leg.; MCZ.

Paratype of Plega fratercula.

USA • 1 ♀; Arizona, Capitan Mt.; 24 Jul. 1933; R. Anderson leg.; “Allotype Plega fratercula Rehn”; MCZ-ENT 00580916, MCZ type 25377; MCZ.

Paratypes of Plega dactylota.

USA • 1 ♂; Arizona, Huachuca Mtn. Carr. Cn.; 06 Aug. 1924; J.O. Martin leg.; “Plega dactylota dactylota Rehn, Paratype”; CAS. • 1 ♂; Huachuca Mts.; 20 Jul. 1937; D.J. & J.N. Knull leg.; Paratype 25379; MCZ. • 1 ♂; Santa Rita Mts.; Jul.; 5–8000 ft; F.H. Snow leg.; Paratype 23379; MCZ. • 1 ♂; Boya Thomson, Aboretum, Superior; 1–3 Aug. 1937; MCZ Paratype 25378; MCZ. – Texas • 1 ♂; Hills W. of Ord Mts Brewster Co.; 22–31 Aug. 1926; O.C. Polling leg.; Paratype 25380, Plega dactylota lipanica; MCZ. • 1 ♂; Hills W. of Ord Mts Brewster Co.; 15–22 Aug. 1926; O.C. Polling leg.; Paratype 25380 Plega dactylota lipanica; MCZ. • 1 ♂; Big Bend Park, Brewster Co.; 14 Jun. 1937; R.H. Baker leg.; Paratype 25380, Plega dactylota lipanica; MCZ. • 1 ♂; Big Bend Park, Brewster Co.; 13 Jun. 1937; R.H. Baker leg.; Paratype 25380, Plega dactylota lipanica; MCZ.

Other material

Mexico • 1 ♀; Chihuahua, 2 mi W. Matachic; 30 Jul. 1984; D. Thomas leg.; TAMU-ENTOX0285142; TAMUIC. – Guerrero • 1 ♀ 1 ?; Grutas de Cacahuamilpa; 02 Nov. 1967; R.E. Woodruff leg.; Blacklight trap; FSCA. – Jalisco • 1 ♀; San Buenaventura; 19°97'61.9"N, 104°03'32.4"W; 720 m; 03 Dec. 1996; S. Zaragoza; CNIN. • 1 ♂; San Buenaventura; 19°47.61'N, 104°03.32'W; 720 m; 07 Nov. 1996; light trap; CNIN. • 1 ♀; Chamela; 12 Jan. 1995; H. Brailovsky; CNIN. • 1 ♀; Estación de Biología Chamela; 24 Oct. 1985; R.A. Usela; trampa de luz; CNIN. • 1 ?; Río Cuitzmala; 24 Nov. 1984; E. Ramírez, J. Villa leg.; CNIN. • 2 ♀; El Limón, San Buenaventura; 620 m; 19°46.614'N, 104°03.324'W; 04 Nov. 1996; S. Zaragoza, F. Noguera, E. Ramírez, E. González leg.; 55037; CNIN. – Morelos • 1 ♂; Tlaquiltenango, 2.5 Km, 4 Km W Huautla Estación CEAMISH; 18°27'67.1"N, 99°02'47.5"W; 940 m; 16 Nov. 1995; S. Zaragoza, F. Noguera, E. Ramírez, E. González leg.; light trap 3, MDMD125; CNIN. • 1 ♂; Tlaquiltenango, Huautla; 18°23'04.27"N, 99°03'00.03"W; 1028 m; 18 Sep. 2009; V.H. Toledo, J.G. Martínez, E.V. Reza, Y.T. Viveros, N. Hernández leg.; light trap, selva baja caducifolia, SHM224; CNIN. • 1 ♂; Tlaquiltenango, Huautla; 18°22'45.01"N, 99°02'44.91"W; 1001 m; 20 Nov. 2009; V.H. Toledo, J.G. Martínez, E.V. Reza, Y.T. Viveros, N. Hernández leg.; selva baja caducifolia, light trap, SHM215; CNIN. • 1 ♀; Tlaquiltenango, Huaxtla; 18°23'8.736"N, 99°3'4.0068"W; 1023 m; 28~31 Jan. 2009; V.H. Toledo leg.; selva baja caducifolia, light trap, SHM113; CNIN. • 1 ♀; Tlaquiltenango, 3 Km E. Santiopan; 18°25'45.66"N, 98°55'56.136"W; 1381 m; 28 Jan. 2014; V.H. Toledo, R. Reyes, J. Martínez leg.; selva baja caducifolia, light trap, SHM124; CNIN. • 1 ♀; El Limón, atrás de estación; 14 Aug. 2015; C. Pérez, P. Bogadrín, G. Gutiérrez leg.; trampa pit light, luz blanca; CNIN. – Nayarit • 1 ♀; P.H. Aguamilpa, Arroyo de los Bueyes; 29~30 Oct. 1991; R. Barba & E. Barrera leg.; CNIN. – Nuevo León • 1 ♂; Bustamante, at entrance of Grutas del Palmito; 08 Oct. 1966; D. Riley; TAMU-ENTOX0070934; TAMUIC. • 1 ♂; Nuevo Leon, nr Monterrey, Mesa de Chipinque; 1365 m; 16–18 Jul.; “Cornell Univ. Mexico Field Party 1965”; MCZ-ENT00681790; MCZ. – Puebla • 1 ♂; Jolalpan, Rancho El Salado; 18°20'49.38"N, 98°59'23.02"W; 923 m; 06 Oct. 2010; V.H. Toledo, F. Hinterholzer, J.G. Martínez leg.; selva baja caducifolia, light trap, SHM221; CNIN. • 1 ♀; Jolalpan, Rancho El Salado; 18°20'49.38"N, 98°59'23.028"W; 923 m; 06 Oct. 2010; V.H. Toledo, F. Hinterholzer, J.G. Martínez leg.; selva baja caducifolia, light trap, SHM178; CNIN. – Querétaro • 1 ♂; Parque Nacional La Joya, La Barreta (cerca a la Monja); 220 m; 06 Aug. 2013; H. Brailovsky, E. Barrera leg.; CNIN. – Sonora • 1 ♂; Nacozari de García, 15.9 Km SE. Nacozari de García, Rancho La Zulema, Sierra Juriquipa; 30°17'2.04"N, 109°33'37.08"W; 1987 m; 15 Jul. 2017; Van Devender, J. Palting leg.; rocky mountainside, oak Woodland; USNMENT001541895; USNM. • 2 ♀ 1 ?; Nacozari de García, 15.9 Km SE. Nacozari de García, Rancho La Zulema, Sierra Juriquipa; 30°17'2.04"N, 109°33'37.08"W; 1687 m; 15 Jul. 2017; Van Devender, J. Palting, leg.; rocky mountainside, oak Woodland; CNIN. • 9 ♂ 12 ♀ 2 ?; Nacozari de García, 15.9 Km SE. Nacozari de García, Rancho La Zulema, Sierra Juriquipa; 30°17'2.04"N, 109°33'37.08"W; 1687 m; 14 Aug. 2017; Van Devender, J. Palting leg.; rocky mountain side, oak Woodland; Plega dactylota Rehn, det. O.S. Flint, 2017; CNIN. • 3 ♀; same data as for preceding; 15 Jul. 2017; CNIN. • 2 ♂ 6 ♀; same data as for preceding; 30°17'45.24"N, 109°36'44.64"W; 14 Jul. 2017; CNIN. • 2 ♂ 2 ♀; Nacozari de García, 15.9 Km SE. Nacozari de García, Sierra Juriquipa; 30°17'45.24"N, 109°36'44.64"W; 1687 m; 14 Jul. 2017; Van Devender, J. Palting leg.; rocky montain side, oak Woodland; CNIN. • 1 ♂; same data as for preceding; Plega dactylota Rehn, det. O.S. Flint, 2017; CNIN. • 2 ♀; Nacozari de García, 15.9 Km SE. Nacozari de García, Sierra Juriquipa; 30°17'1.68"N, 109°40'9.48"W; 1687 m; 15 Jul. 2017; Van Devender, J. Palting leg.; rocky mountain side, oak Woodland; CNIN. • 1 ♀; Nacozari de García, Pilares de Nacozari, 6.5 Km (by air) SE of Nacozari de García, Sierra Nacozari; 30°19'41.988"N, 109°37'46.992"W; 1413 m; 09 Aug. 2015; T.R. Van Devender, A.L. Reina-G leg.; rocky slope; CNIN. • 4 ♂ 3 ♀; Nacozari, rancho El Metate, 7.4 Km SSW Nacozari, E. base of Sierra San José; 31°15'30.96"N, 109°57'33.12"W; 1634 m; 16 Sep. 2017; Van Devender, J. Palting leg.; desert grassland, oak Woodland; CNIN. • 1 ♀; Sonora, Agua Caliente; 25 Aug. 1959; R.E. Ryckman, C.P. Christianson & D. Spencer leg; USNM. • 2 ♂ 4 ♀; Mpio. Álamos, Parque La Colorada, sendero Arroyo alto (Chalatón); 27°00.551'N, 108°57.206'W; 510 m; 17 Sep. 2019; Contreras, Barba, Cancino, Ardila, Marquez leg.; selva baja caducifolia, black and white light trap; CNIN. • 4 ♂ 1 ♀; Mpio. Álamos, Parque La Colorada, sendero Tecolote; 27°00.675'N, 108°56.987'W; 469 m; 16 Sep. 2019; Contreras, Barba, Cancino, Ardila, Marquez leg.; selva baja caducifolia, black and white light trap; CNIN. • 2 ♂; Mpio. Moctezuma, Rancho La Montosa, camino; 29°34.201'N, 109°58.352'W; 930 m; 22 Nov. 2019; Barba, Ardila, Cancino, Marquez, Contreras leg.; black and White light trap, bosque espinoso; CNIN. • 1 ♀; 36.6 Km SE Tecoripa, La Barranca; 28°34'40.1"N, 109°39'54.3"W; 562 m; 11 Sep. 2004; S. Zaragoza, F. Noguera, E. Ramírez, E. González leg.; light trap 1, bosque tropical caducifolio, MDM0233, 2699; CNIN. • 1 ♀; same data as for preceding; MDM0243, 2720; CNIN. • 1 ♀; same data as for preceding; MDM0241, 2719; CNIN. • 1 ♂; same data as for preceding; 28°34'40.1"N, 109°39'54.3"W; 645 m; 16 Aug. 2004; MDM0262, 2722; CNIN. • 1 ♀; same data as for preceding; 28°34'40.1"N, 109°39'54.3"W; 645 m; 16 Aug. 2004; MDM0260, 2717; CNIN. • 3 ♂ 3 ♀; Rancho Los Jarazos, Sierra La Purica; 16 Jul. 2013; T.R. Van Devender, J. Palting leg.; CNIN. – Tamaulipas • 1 ♀; 43 km. east 27.V Ciudad Mante; FSCA.

USA – Arizona • 1 ♂ 1 ♀; Cochise Co., S.W. Research Sta., nr. Portal; 1982; L.L. Lampert leg.; black light trap; FSCA. • 1 ♂; Chochise County, SW Research Sta.; 27 Jul. 1989; A.M. & N.D. Penny leg.; at lights; Plega dactylota Rehn, det. N. Penny; CAS. • 1 ♂ 1 ♀; Cochise Co., Huachuca Mtns., Ash Canyon; 07 Aug. 1991; R. & J. Robertson leg.; Plega dactylota, det. Penny, 1991; CAS. • 2 ♀; same data as for preceding; 22 Jul. 1989; CAS. • 1 ♂; Cochise Co., Huachuca Mts., Ash Canyon; 5100’; 09 Aug. 2001; B.D. Valentine leg.; Plega dactylota Rehn, det. D.J. Shetlar, 1994 and Penny, 1997; CAS. • 1 ♂; Cochise Co., Huachuca Mtns., Ash Canyon; 27–31 Jul. 1986; D.L. Wagner leg.; black and white lights; CAS. • 2 ♂ 5 ♀; Cochise Co., Ramsey Cyn.; 31°26'57.84"N, 110°18'23.40"W; 1679 m; 22–24 Jul. 2017; Aalbu R.L. leg.; black light; CSCA. • 1 ♀; Cochise Co., 3.2 mi E. SERS; 16 Aug. 1971; D. Carlson leg.; CSCA. • 1 ♀; Gila Co., Carrizo; 4750’; 01 Sep. 1986; A.S. Menke leg.; USNM. • 1 ♂; Portal; 10 Jul. 1969; G.H. Nelson Family leg.; at lite; FSCA. • 1 ♂ Gila Co., Payson; 30 Aug. 2001; P.R. Coleman leg.; UCD. • 1 ♂; Maricopa Co., Jct. Fish, Ck. Cyn & Apache Trail; 01 Aug. 1987; W.B. Warner; U.V. light; FSCA. • 1 ♂; Santa Rita Mts., Madera Canyon; 14 Aug. 1949; 00045; “Phillip A. Adams Collection 1998 request to Calif. Acad. Sci.”; CAS. • 1 ♂; Santa Cruz Co., Duquesne Rd. (For. Ser. Rd. #61), 7.8 mi. E. Jct. Hwy. 82; 4800 ft.; 13 Aug. 1991; L.G. Bezark, D.E. Russell, R.A. Cunningham leg.; MV/BL, Oak/Mesq. Trans; Plega fratercula, det. N. Penny, 1997; CAS. • 1 ♀; Santa Cruz Co., Sta. Rita Mts., Madera Canyon, Sta. Rita Lodge; 27 Aug. 1991; R. Wielgus leg.; CAS. • 1 ♂; Santa Cruz Co., Madera Cyn.; 4880’; 15 Jun. 1965; D.N. Harrington leg.; UCD. • 3 ♂; same data as for preceding; 27 Jul. 1965; D.N. Harrington leg.; UCD. • 3 ♂; same data as for preceding; 07 Sep. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 26 Aug. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 21 Aug. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 20 Aug. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 15 Aug. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 07 Sep. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 15 Sep. 1965; D.N. Harrington; UCD. • 1 ♂; same data as for preceding; 01 Oct. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding; 07 Sep. 1965; D.N. Harrington leg.; UCD. 1 ♂; same data as for preceding; 01 Oct. 1965; D.N. Harrington leg.; UCD. • 1 ♂; same data as for preceding: 17 Jul. 1963; V.L. Vesterby; UCD. 1 ♂; same data as for preceding;17 Jul. 1963; V.L. Vesterby; UCD. • 2 ♂; same data as for preceding; 21 Jul. 1963; V.L. Vesterby; UCD. • 5 ♂; same data as for preceding; 15 Aug. 1963; V.L. Vesterby; UCD. • 3 ♂; same data as for preceding; 12 Aug. 1963; V.L. Vesterby; UCD. • 3 ♂; same data as for preceding; 25 Aug. 1963; V.L. Vesterby; UCD. • 1 ♂; same data as for preceding; 08 Sep. 1963; V.L. Vesterby; UCD. • 1 ♂; same data as for preceding; 08 Jul. 1963; V.L. Vesterby; UCD. • 1 ♂; same data as for preceding; 22 Sep. 1963; V.L. Vesterby; UCD. • 1 ♂; same data as for preceding; 12 Sep. 1963; V.L. Vesterby; UCD. • 1 ♂; Maricopa Co., Sunflower, Jct. Hwy, 87 & Sycamore Ck.; 33°52'8.616"N, 111°27'51.732"W ± 100 m; 744 m; 15 Aug. 2012; Oswald, Diehl, Machado leg.; MV light; TAMU-ENTO X0965263; TAMUIC. • 1 ♀; same data as for preceding; TAMU-ENTO X0966388; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTO X0966101; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTO X0966424; TAMUIC. • 1 ♂; Pima Co., Madera Cyn., Santa Rita Mtns.; 4400 ft.; 1–2. Aug. 1975; Menke & Pulawsky leg.; CAS. • 2 ♂; Pima Co., Santa Rita Mts Experimental Range, Florida Cyn.; 31°45.42'N, 110°50.43'W; 1646 m; 03–10 Jul. 2014; M.I. Erwin, C. O’ Brien, M.J. Sharkey leg.; Malaise on hillside road; CSCA. • 1 ♀; same data as for preceding; 23 Aug. 2014; CSCA. • 2 ♀; same data as for preceding; 10–20 Sep. 2014; CSCA. • 1 ♂; Pima Co., Sabino Cyn.; 03 Sep. 1962; V.L. Vesterby; UCD. • 1 ♂; same data as for preceding; 04 Sep. 1961; UCD. • 1 ♂; same data as for preceding; 14 Sep. 1963; UCD. • 1 ♂; Organ Pipe Cactus Nat. Mon.; 13 Oct. 1963; V.L. Vesterby; UCD. • 1 ♀; Mohave Co., 11 Km E. Kingman on Hualapai Mt. Rd.; 35°58.55'N, 113°55.63'W; 1490 m; 19 Jul–01 Aug. 2018; M.E Irwin leg.; Malaise in narrow wash nr Hualapai Mt. Recreational Park; CSCA. • 1 ♂ 1 ♀; Santa Cruz Co. Sonoita, 3.2 Km Town Center; 31°38'N, 110°39'W; 1524 m; 21 Jul.–06 Aug. 2014; E.E. Grissell leg.; Malaise in juniper-oak-grass land; CSCA. • 1 ♂ 1 ♀; Gila Co., W. side of Pinal Mtn.; 14 Aug. 1977; R.P. Allen leg.; black light; CSCA. • 1 ♂; Gila Co., 5 mi N Payson; 05 Sep. 1961; J.S. Bucket leg.; UCD. • 1 ♂; same data as for preceding; 05 Sep. 1961; UCD. • 1 ♀; 5 mi. Portal Chiricahua Mts.; 17 Aug.1958; G.B. Pitman leg.; UCD. • 1 ♂; 15 mi. W. Portal Chiricahua Mts.; 04 Aug. 1958; R.E. Rice; UCD. – New Mexico • 1 ♂; Grat Co., Hwy. #15, 5.5 Km S Jct. #35 3258/10812; 2165 m; 22 Jul. 1983; E.L. & K.W. Sleeper, B.L. leg.; Pine Oak Fst Mdw; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; Catron Co., Gila N.F., Whitewater Pub. Camp.; 11–14 Jul. 1970; J.A. Gruwell leg.; Plega dactylota, det. N. Penny, 1998; collected at blacklite; CAS. • 1 ♀; Dona Ana Co., Aguirre Sprgs RA; 5600’; 26–27 Aug. 1997; Wappes & Turnbow leg.; TAMU-ENTOX0070958; TAMUIC. • 1 ♀; same data as for preceding; TAMU-ENTOX0070968; TAMUIC. • 1 ♀; same data as for preceding; TAMU-ENTOX0070972; TAMUIC. • 8 ♂; Lincoln Co., HWY 7018-50 Km E Ruidoso; 33°22.1'N, 105°29.9'W; 01 Jul. 2013; C.M. Parker leg.; funnel light trap; UCD. • 10 ♂; same data as for preceding; 01 Jun. 2013; UCD. – Texas • 1 ♂; Jeff Davis Co., Davis Mountains Resort; 30°37'42"N, 104°05'01"W; 5800 ft.; 17 May. 2001; D.G. Marqua leg.; FSCA. • 1 ♀; Jeff Davis Co., Davis Mtns. Resort; 5800’; 31 Aug. 1999; D.G. Marqua leg.; TAMU-ENTOX0085828; TAMUIC. • 1 ♂; Jeff Davis Co. rest stop, 9.5 mi. S. jct. hwy 118 on 166; 09 Aug. 1992; W. Godwin & E. Riley leg.; UV light; TAMU-ENTOX0070888; TAMUIC. • 1 ♂; Jeff Davis Co., Davis Mountains Resort; 30°37'42"N, 104°05'01"W; 13 Aug. 2001; 5800 ft.; D. Marqua leg.; UV; UCD. • 1 ♀; same data as for preceding; 15 Aug. 2001; UCD. • 1 ?; Brewster Co., Big Bend Park; 26 Jul. 1937; R.H. Baker; TAMU-ENTOX0070916; TAMUIC. • 1 ?; same data as for preceding; 15 Jul. 1937; TAMU-ENTOX0071037; TAMUIC. • 1 ♂; Brewster Co., BBNP, Basin Area; 29°16'12"N, 103°18'01"W; 18–19 Jul. 2002; E.G. & C.M. Riley leg.; at lights; Plega dactylota Rehn, 1930, det. J.D. Oswald, 2004; TAMU-ENTOX0281148; TAMUIC. • 1 ♀; same data as for preceding; TAMU-ENTOX0281153; TAMUIC. • 1 ♂; Brewster Co., Big Bend National Park, Basin Lodge; 5400 ft.; 08 Jul. 2001; D.G. Marqua leg.; Plega dactylota Rehn, 1930, det. J.D. Oswald, 2002; TAMU-ENTOX0088735; TAMUIC. – Utah • 1 ♂; Juab Co., Hwy. 132 E. of Nephi, 1.5 mi. S. of Nebo Loop Rd.; 21 Mar. 1998; R.C. Mower leg.; Plega dactylota Rehn, 1930, de J.D. Oswald, 2003; TAMU-ENTOX0090578; TAMUIC. • 1 ♀; Juab Co., Hwy. 132 E. of Nephi, 1.5 mi. S. of Nebo Loop Rd.; 21 Aug. 1998; R.C. Mower leg.; TAMU-ENTOX0090575; TAMUIC.

Diagnosis

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.

Description

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. 32). Head. Pale with dark brown marks; vertexal pale with lateral, parallel, dark brown bands extended from occipital suture to supra-antennal area and enclosing lateral pale areas at level of paraocular region; area adjacent to occipital ridge dark brown; area adjacent to coronal suture pale to brown, sometimes arrow-shaped; supra-antennal area dark brown; brown setae present; occiput pale with dark brown mark, postgena pale with brown area. Antennal scape brown with pale outer surface, pedicel dark brown; flagellum pale to dark brown. Frons dark brown, sometimes with medial, pale area. Clypeus dark brown with pale anterior margin. Labrum brown; mandible pale with dark brown corners, dark amber at apex; maxilla pale with dark brown areas, palpus brown; labium pale with brown postmentum, ligula and palpus; palpimacula pale brown. Thorax. Pronotum pale with broad, brown, medial band, anterolateral areas dark brown; episternum brown; postfurcasternum pale. Mesonotum mostly dark brown, often with pale areas adjacent to sutures; scutellum sometimes with pale medial area. Metanotum dark brown, sometimes with pale anteromedian area; pre-episternum brown; pteropleura dark brown with pale areas. Foreleg. Coxa from pale with brown areas, to pale or dark brown or reddish brown; with interspersed pale and dark brown setae, sometimes mostly dark brown, setal bases pale to dark brown; trochanter pale with brown suffusions to brown with paler areas, setation as in coxa. Femur posterior surface either nearly completely brown or pale with two pairs of brown areas near medial region of femur; apex of posterior surface brown; anterior surface from brown to dark reddish brown, with pale proximal part of closing surface. Tibia posterior surface alternating pale and brown, anterior surface mostly dark brown, with preapical pale area. Basitarsus with base and lanceolate process dark amber, sub-basal area pale brown area, clavate setae brown; remaining tarsomeres pale brown. Mid- and hind leg. Coxae brown with paler areas; trochanter pale with brown areas. Femora and tibiae dashed, alternating pale and brown. Tarsi pale, becoming pale brown, towards the last tarsomere. Wings. Forewing mostly hyaline; membrane surrounding crossveins, R fork, stem of RP branches, M fork, first branch of CuA and apex of CuP, and terminal forks of longitudinal veins amber. Pterostigma brown with pale medial area. Major veins, subcostal veinlets and wing margin alternating pale and brown; crossveins dark brown. Hind wing hyaline, amber on area adjacent to stem first branch of CuA; pterostigma brown with pale preapical area; longitudinal veins mostly brown, with small pale areas; subcostal veinlets pale; crossveins brown, 1r-m bicolor; wing margin alternating pale and brown. Abdomen. Tergites dark brown. Pleural membrane dark brown with pale areas. Sternites brown, with paler medial and lateral areas.

Figure 32. 

Plega dactylota Rehn, 1939 A female habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 32). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct; medial area of ​​supra-antennal region not raised, with fine, reclined setae; paraocular area concave. Antenna sub-moniliform; scape as two times as long as wide, ventrally with a few thickened setae; pedicel as long as wide; flagellum slightly dorsoventrally flattened, with 42–56 flagellomeres, those of proximal ¼ discoidal, then changing from two times as long as wide to as long as wide; all articles with medial ring of fine, short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus narrow. Labrum pentagonal; maxillary palpus with first palpomere as long as wide, second 1.5× as long as wide, third palpomere 4× as long as wide, fourth palpomere 4× as long as wide, fifth palpomere as long as third; mentum with long, thin setae; labial palpus with first palpomere 2.5× as long as wide, second palpomere 5× as long as wide, third palpomere slightly longer than second, palpimacula ovoid. Thorax. Pronotum as long as wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae. Mesonotum slightly wider than long, with abundant, thick, pedicellate setae on medial area. Metanotum ~ 2× as wide as long. Pteropleura with short, thin setae, thicker on mesepisternum and metakatepisternum. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with pedicellate, fine or thick setae of different sizes; trochanter trapezoidal, dorsal surface with some thickened, pedicellate setae near distal margin; anterior surface without protuberant area. Femur robust, closing surface with posteroventral row of processes composed two medially located, primary processes; proximal portion of the row with sub-basal secondary process and basal tertiary process; the rest of the row with abundant tubercle-shaped processes, and stinger-shaped setae; distal portion carinate, composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½; composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ⅘. Tibia almost as long as femur, curved, ventral surface keeled with row of prostrate setae; patch of clavate setae apically on anterior surface present. Basitarsus with lanceolate process reaching base of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere about 7× as long as wide; third tarsomere as long as wide; fourth tarsomere ~ 3× as long as wide. Mid- and hind leg. Coxa and trochanter with thin setae, shorter on trochanter; femora with interspersed fine setae of different lengths and few thickened setae; tibiae mostly with short fine setae; hind leg longer than midleg, tibia 1.5× as long as femur; tarsomeres with distal margin of plantar surface with lateral groups of 5–8 thickened setae; on both legs, basitarsus 4× as long as wide, second tarsomere 1.2× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing narrowly oval, trichosors present along margin except on wing base; costal space proximally expanded, humeral vein branched; 14–18 subcostal veinlets; pterostigma elongated, narrow, straight, with incomplete veinlets; subcostal space with single, medially located crossvein; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 gently curved with three or four RP branches; four veins arising from rarp1; M vein basally fused with RA; RP base widely separated from divergence of M and R; M forked opposite to RP origin; 1 r-m connecting RP base and M fork, forming a trapezoidal cell; 5–7 gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked oppositely to level of separation of M and R; A1 apically forked, ending on posterior margin at level of CuP fork, A2 forked oppositely to CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with 6–8 veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge RA; pterostigma elongated, narrow, gently curved, composed of incomplete veinlets; radial space with single crossvein, oblique; four veins arising from rarp1, one or two from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent or present; CuP strongly anteriorly bent at distal ½, pectinate; two crossveins on cubitoanal space; A1 apically simple, ending on wing margin at 1r-m stem level; A2 generally simple, short. Abdomen. Cylindrical to medially expanded, with subquadrate tergites; tergites III and IV with posteromedian concavity; tergites III-VII with elongated anterolateral scars; sternites rectangular.

Male genitalia (Fig. 33A–E). Tergite IX medially narrower than laterally; lateral margin notched. Sternite VIII rectangular; sternite IX pentagonal in ventral view; posterior margin with short, medial, narrow lobe which is dorsally canaliculated; in lateral view triangular, apex not surpassing posterior margin of ectoproct. Gonocoxites IX elongate, thickened, sinusoid; base flattened, spatulated; apex branched, with 5–15 closely adpressed, twisted processes. Ectoproct ovoid, anteroventrally with rounded, flattened lobe that is continuous with ventromedial sclerotized, curved sulcus. Gonocoxites X forming a short, ventrally canaliculated sclerite, whose posterior apex has dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X; gonostyli X with thickened and curved base, equipped with two lateral processes, the rest of the structure, ventrally curved, and anteriorly coiled, forming two loops before protruding from abdomen. Gonapophyses X long, straight, and narrow, with dorsally curved apexes; gonapophyses arranged in a V-shaped structure, joined by a membrane covering the gonostyli X base. Gonocoxites XI U-shaped, medial lobe complex and elaborated, with two differentiated parts: dorsal part as an expanded arch which has less sclerotized, transparent, medial area from which a ventrally incised process protrudes; ventral part as a curved, short lobe; lateral arms of gonocoxites XI straight, with anterior apex curved posteroventrally. Hypandrium arched and concave.

Figure 33. 

Plega dactylota Rehn, 1939 A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 33F–I). Sternite VII medially narrower than laterally, posterior margin concave, with long, thickened setae. Tergite VIII narrower medially than laterally, encircling the spiracle of the segment; lateral margin bluntly quadrangular. Gonocoxites + gonapophyses VIII medial part keel-shaped, posteromedially produced into a short bilobed process; lateral part an oval plate. Tergite IX + ectoproct ovoid. Gonocoxites IX elongated, sinuous, and narrow, as long as the last seven abdominal segments together. Bursa copulatrix short, conical, and moderately sclerotized. Spermatheca long, complex and entangled; proximal section thin, long, forming several coils; medial section as wide as proximal section, forming a long spiral; distal section forming several convolutions, distally widened, forming a diverticulum; fertilization canal duct proximally triangular, concave; fertilization canal elongated, curved, covered with microfilaments.

Distribution

Mexico (Chihuahua, Coahuila, Guerrero, Jalisco, Morelos, Nayarit, Nuevo León, Puebla, Querétaro, Sinaloa, Sonora, Tamaulipas), USA (Arizona, Colorado?, New Mexico, Texas, Utah).

Remarks

This species is widely distributed in Mexico and the United States. The records from Coahuila and Sinaloa were extracted from Oswald et al. (2002) and Reynoso-Velasco and Contreras-Ramos (2008). The records from Baja California Sur provided by these same authors are more likely erroneous and based on the material studied herein, such reports correspond to P. flammata. Here, this species is newly recorded for the Mexican states of Guerrero, Morelos, Nayarit, Nuevo Leon, Puebla, and Tamaulipas. Rehn (1939b) reported this species from Arizona (Cochise and Mohave Counties), Nevada (Charleston), Utah (Stockton and Duchesne) and Texas (Brewster and Jeff Davis Counties). The records from California provided by Penny et al. (1997) are likely doubtful and these probably represent specimens of P. signata. Here, this species is newly recorded from New Mexico.

This species is highly variable in coloration, size, and morphology. Rice (1987) studied the variation on the number of processes of the male gonocoxites IX, reporting from seven to twelve processes, leading to the synonymy of Plega dactylota lipanica Rehn. Based on the examination of the type specimen of P. fratercula and numerous specimens of P. dactylota, it was concluded the former represent a synonym of the latter. In the original description by Rehn (1939b), this species was separated from other North American species based on its body size and the presence of five processes on the gonocoxites IX. The overall arrangement of these processes was also used to distinguish this species, but this condition is noticeably similar to that expressed by P. dactylota. In this work, several small- and large sized specimens with only four processes were studied, leading to the conclusion that this character is quite plastic, and the variation is even broader to that reported by Rice (1987). Additional support for this synonymy is based on the morphology of the remaining genital sclerites of the male of P. fratercula being identical to those of P. dactylota. Finally, the distribution of P. fratercula from a single locality in Arizona is entirely overlapped with that of P. dactylota which is wide in this state.

Plega disrupta Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 34, 35

Type locality

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”.

Material examined

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”, CNIN. Paratypes. Mexico • 1 ♀; same. data as for holotype; 12 Dec. 2016; CNIN.

Other material

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”; CNIN. • 1 ♂; same data as for preceding; 12 Jan. 2011; CNIN.

Diagnosis

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.

Description

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. 34). Head. Vertexal region pale with V-shaped inverted mark extended from occiput to supra-antennal region, with pale brown setae. Occiput with dark brown marks near occipital transverse suture, setation pale brown; postgena pale with enlarged dark brown mark. Supra-antennal region mostly dark brown. Antennal scape pale with dark brown, dorsal, longitudinal band, and ventral mark, with dark brown near apex; pedicel dark brown, flagellum brown, with single paler preapical flagellomere. Frons mostly dark brown with medial area pale. Clypeus pale with lateral dark brown marks; labrum pale; mandible pale with dark brown dorsal and ventral corners, apex brown; maxilla pale with dark brown marks, palpus dark brown, except for pale fourth palpomere, and junctions of the remaining palpomeres, setae brown; labium pale with dark areas, setae brown; labial palpus dark brown, with pale junctions, palpimacula pale brown. Thorax. Pronotum mostly brown with anterolateral and posterolateral small, pale areas, and dark lateral margins; setae mostly dark brown and a few pale brown; episternum mostly dark brown with brown setae; postfurcasternum pale with lateral dark brown area. Mesonotum mostly dark brown with small pale areas, setation dark brown; metanotum dark brown with paler anteromedial area; pre-episternum dark brown mark; pteropleura dark brown with pale areas, with interspersed dark and pale brown setae. Foreleg. Coxa pale with basal and apical brown marks which are connected through brown suffusions on anterior and posterior surfaces; setal bases dark brown; setae mostly dark brown; trochanter pale with dark brown marks, interspersed pale and dark brown setae present. Femur posterior surface pale with dark brown dots, apex dark brown; anterior surface dark brown; dorsal surface with some suffusions connecting anterior and posterior maculations. Tibia dashed with pale and dark brown. Basitarsus with base and lanceolate process amber, proximal ½ with pale area, clavate setae pale brown; remaining tarsomeres pale with brown areas, setae brown; pretarsal claws pale amber. Mid- and hind leg. Coxae pale with brown areas, setation mostly pale brown; trochanter pale with dark brown marks. Femora and tibiae pale with dark brown rings. Tarsi pale, except the fifth tarsomere pale brown, setae dark brown; pretarsal claws pale amber. Wings. Forewing mostly hyaline or slightly amber shaded, dark amber on area surrounding crossveins, and intermittent amber areas on apical twigging near posterior and apical wing margin; major veins and subcostal veinlets alternating pale and dark brown, crossveins dark brown; pterostigma brown with pale, sub-basal area; wing margin alternating pale and brown. Hind wing mostly hyaline, with intermittent amber infuscation on the apical twigging of major veins near the posterior and distal margins; pterostigma dark brown with preapical pale area, and darker marks on anterior margin; longitudinal veins alternating pale and brown, except CuA mostly dark brown; subcostal veinlets and crossveins dark brown; wing margin alternating brown and pale. Abdomen. Tergites dark brown with pale lateral areas, setae mostly pale brown. Pleural membrane dark brown. Sternites pale with lateral, subtrapezoidal brown marks.

Figure 34. 

Plega disrupta Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 34). Head. Diamond-shaped in frontal view, rugose, vertexal region raised above compound eyes, with lateral rows of reclined setae; area adjacent to coronal suture and occipital ridge, glabrous, with muscle insertion mark; coronal suture distinct, ​​supra-antennal region slightly raised, covered with reclined setae; paraocular area concave. Antenna filiform, short; scape 1.5× as long as wide, slightly distally expanded, with short setae, a few longer and thicker near distal margin; pedicel as long as wide; flagellum not dorsoventrally flattened, with 30–33 or 41 flagellomeres, which are as wide as long on proximal part, and longer than wide on the rest of the flagellum; all articles with medial ring of short setae. Compound eye hemispherical, as wide as ¼ of the interocular distance at torulus level. Frons and clypeus rugose, with fine and short setae. Labrum pentagonal with fine and short setae; maxillary palpus with first palpomere as long as wide, second 1.5× as long as wide, third palpomere 3× as long as wide, fourth palpomere two times as long as wide, fifth palpomere slightly 1.5× as long as third, all the articles with short and thin setae; mentum with thin, long setae. Labial palpus with first palpomere two times as long as wide, second palpomere 4× as long as wide, third palpomere slightly longer than second, palpimacula ovoid. Thorax. Pronotum slightly longer than wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae, remaining surface with fine, short setae; postfurcasternum ventrally fused, collar-like. Mesonotum 2× as wide as long, with scattered, thick, pedicellate setae and some long, thin setae on medial area. Metanotum ~ 2× as long as wide, mostly glabrous with few fine setae on scutellum. Pteropleura with short, thin setae, and a few thick, longer setae. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with pedicellate, fine or thick setae of different lengths; trochanter subconical, with thin and short setae, except on dorsal and anterior surfaces with some pedicellate, thick, long setae. Femur robust, covered with abundant, fine, short setae which arise from protuberant bases on both surfaces. Closing surface with posteroventral row of processes composed two medially located primary processes and two sub-basal secondary process; between these processes a single tertiary process, and abundant tubercle-shaped processes and stinger-shaped setae are preset; distal portion of posteroventral row raised, composed of proximal tertiary process, tubercle-shaped specializations of different sizes and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½ and distal region; it is composed of tubercle-shaped specializations, stinger-shaped setae, and a basal primary, curved process; distal portion composed tubercle-shaped specializations; adjacent row of thickened setae with globular base present on distal ¾. Tibia almost as long as the femur, curved, setose, ventrally keeled with prostrate setae row; a patch of clavate setae is present apically on the anterior surface. Basitarsus with lanceolate process reaching the middle of fourth tarsomere; clavate setae are present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 7× as long as wide; third tarsomere slightly longer than wide, fourth tarsomere 3× as long as wide. Mid- and hind leg. Coxae with thin, long setae; trochanter with fine and short setae; femora and tibiae with thin setae of different lengths; tibial spurs short; hind leg longer than midleg, tibia, 1.5× as long as femur; tarsi with thin, short setae, except on distal margin of plantar surface with lateral groups of 3–5 thickened setae; on both legs basitarsus 4× as long as wide, second tarsomere 1.5× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing oval, trichosors present along margin except on wing base; venation setose; costal space proximally expanded, humeral vein branched, 14–17 subcostal veinlets, sometimes with a few apical forked; pterostigma elongated, trapezoidal, with well-defined, mostly forked, entire veinlets; subcostal space with single crossvein, medially located; Sc vein abruptly posteriorly curved at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 curved with three RP branches arising from it; four veins arising from rarp1; M vein fused basally with RA; RP base widely separated from point of divergence of M from R; M forked opposite to R fork, 1 r-m connecting RP base and MA stem, forming a trapezoidal cell; six or seven gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked at level of separation of M and R; A1 apically forked, ending on posterior margin at CuP fork level, A2 forked at CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with five or six veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriorly at proximal margin of pterostigma to merge with RA; pterostigma elongated, narrow, composed of entire, mostly forked veinlets; radial space with single crossvein, oblique; three or four veins arising from rarp1, one or two from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent; CuP not touching A1, strongly anteriorly bent at distal 1/3, with three branches; two crossveins on cubitoanal space; A1 simple ending on wing margin slightly beyond 1m-cu level; A2 simple, short and curved. Abdomen. Cylindrical to medially expanded, setae on tergites gradually longer and more abundant towards terminal segments; tergites III–VI with elongated anterolateral scars. Sternites rectangular, with abundant, thin, long setae.

Male genitalia (Fig. 35A–E). Tergite IX medially narrower than laterally, lateral margin bluntly quadrangular. Sternum VIII rectangular, wider medially than laterally; sternite IX trapezoidal in ventral view, with lateral indentation, covered with abundant thin, short setae, posteromedially produced into a triangular lobe, which is equipped with dorsal canal; in lateral view trapezoidal, apex not reaching posterior margin of ectoproct. Gonocoxites IX filiform, thin, sinusoid; base connected to gonocoxites XI with a membrane; apex pointed, without processes. Ectoproct ovoid, with thin and short setae; ventral surface with anterior rounded lobe, which is continuous with sclerotized, incurved, ventromedial sulcus. Gonocoxites X forming a short, ventrally canaliculate sclerite, whose anterior apex is spatulated; posterior apex with dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X with a membrane; gonostyli X with thickened, gently curved, with two lateral processes at base; the rest of the structure, whip-shaped, ventrally curved, and anteriorly recurved, forming two loops before protruding from abdomen. Gonapophyses X as two short, thin, rod-shaped, straight sclerites; the gonapophyses are joined by a membrane which is medially sclerotized and covers the gonostyli X base. Gonocoxites XI U-shaped, medial lobe complex and elaborated, with two differentiated parts: a dorsal arch and a posteroventral U-shaped projection, with lateral rounded fins; between these parts a narrow, arched, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI thin, short, straight.

Figure 35. 

Plega disrupta Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H gonapophyses VIII of female, ventral I spermatheca.

Female genitalia (Fig. 35F–I). Sternum VII rectangular, posteromedially continuous with sclerotized, glabrous area, the rest of the surface with thin and short setae. Tergite VIII slightly narrower medially than laterally, encircling the spiracle of the segment, lateral margin bluntly quadrangular. Gonocoxites VIII as a rounded, concave plate. Gonapophyses VIII as two medially joined plates forming a tubular structure; posteromedially with two lateral, digitiform lobes, and medial U-shaped concavity; median part of gonapophyses VIII connected to narrowly, oval lateral plates, which are folded beneath tergite IX + ectoproct. Tergite IX + ectoproct narrowly ovoid. Gonocoxites IX elongated, sinuous, and narrow, as long as the last five abdominal segments combined. Bursa copulatrix long, funnel-shaped, membranous with area adjacent to genital pore strongly sclerotized. Spermatheca relatively long and complex; proximal section, long, thin, forming numerous coils; medial section thin, as wide as proximal section, forming numerous convolutions; distal section slightly expanded, wider than medial and proximal sections, but abruptly narrowed at apex; fertilization canal duct short, triangular, concave, with distal, thin portion; fertilization canal C-shaped, covered with microfilaments.

Etymology

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.

Distribution

Mexico (Oaxaca).

Remarks

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.

Plega drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 36, 37

Type locality

Mexico, Jalisco: Sierra Manatlán, Puerto Los Mazos, 1500 m, 3–7 Aug. 1992, Luz Hg, Encinar, D. Curoe leg.

Material examined

Holotype male, pinned. Original label: “Mexico, Jalisco, Sierra Manatlán, Puerto Los Mazos, 1500 m, 3–7 Aug. 1992, D. Curoe leg., Luz Hg, Encinar”, CAS. Paratypes. Mexico – Chihuahua • 1 ♂; Rta. San Rafael-Cuiteco; 27°28.130'N, 107°56.580'W; 1870 m; 30 Aug. 2005; J. Bueno, R. Barba leg.; CNIN. – Jalisco • 1 ♂; San Sebastián del Oeste; 20°46'5.04"N, 104°50'27.67"W; 1620 m, 29.VII.2019, A. Ramírez-Ponce Leg., light trap, mixto de pino-encino 2743 (CNIN).

Other material

Mexico • 1 ?; Jalisco, Sierra Manatlán, Puerto Los Mazos; 1500 m; 3–7 Aug. 1992; D. Curoe leg.; Luz Hg, Encinar; CAS. • 1 ♂; Jalisco, Los Mazos; 15 Jul. 1989; R.J. McGinley leg.; USNM. • 1 ♀; Jalisco, La Yerbabuena; 1950 m; 04 Aug. 1994; G. Nogueira; CNIN. • 1 ♂; Volcán de Tequila; 2150 m; 10–11 Aug. 1994; G. Nogueira; CNIN. • 1 ♀; San Sebastián del Oeste; 20°46'5.04"N, 104°50'27.67"W; 1620 m; 29 Jul. 2019; A. Ramírez-Ponce leg.; light trap, mixto de pino-encino; CNIN 2739. • 1 ♂; same data as for preceding; CNIN 2741. • 1 ♂; same data as for preceding; CNIN 2742. • 1 ♀; same data as for preceding; CNIN 2740. • 1 ♂; same data as for preceding; CNIN 2744. • 1 ♂; Volcán de Tequila, 3.7 km S Hwy 15d; 20°49'27"N, 103°51'25"W; 1849 m; 09 Jul. 2017; E. Martínez leg.; CSCA.

Diagnosis

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.

Etymology

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.

Description

Measurements. Male (n = 4). Forewing length: 11.9–15.4 mm; Hind wing length: 9.9–13.2 mm.

Coloration (Fig. 36). Head. Vertexal region pale with dark brown paired irregular markings extending from occiput to toruli, with interspersed pale and dark brown setae. Occiput with dark brown marks near occipital transverse suture, setation brown; postgena pale. Antennal scape pale with dark brown longitudinal ventral and dorsolateral bands, distal margin dark brown, setae dark brown; pedicel dark brown, flagellum brown. Frons pale with dark brown lateral markings which merge anteromedially. Clypeus pale with lateral dark brown markings; labrum pale with anterior ½ brown; mandible pale with dorsal and ventral corners, plus apex dark brown; maxilla pale with dark brown markings, palpus dark brown, except for pale junctions between segments, setae brown; labium pale with pale brown areas, setae dark brown on anterior portion of submentum and anterior area of prementum; labial palpus dark brown, third palpomere paler, palpimacula pale brown. Thorax. Pronotum pale with dark brown lateral stripes, medial area with irregular markings, setae mostly dark brown and a few pale brown; episternum mostly brown with interspersed yellow, pale- and dark brown setae; postfurcasternum pale. Mesonotum mostly dark brown with small anterolateral and medial pale areas, setation dark brown; metanotum dark brown with pale area on medial region of scutum; pre-episternum pale with dark brown mark; pteropleura pale with brown markings. Foreleg. Coxa pale with basal and apical brown markings which are connected through brown suffusions; trochanter pale with dark brown marking on anterior surface, setae of dorsal surface dark brown. Femur posterior surface pale with discontinuous brown spots on proximal and distal ½, apex dark brown; anterior surface mainly dark brown with small pale areas, setal bases bordered with dark brown forming a honey-comb pattern. Tibia dashed with pale and dark brown. Basitarsus with base and lanceolate process amber, proximal ½ with pale area, clavate setae pale brown; second tarsomere brown, third and fourth paler with brown setae. Mid- and hind leg. Coxae pale with small brown areas, pale brown setae present; trochanter pale with dark brown spots. Femora and tibiae pale with brown spots, hind leg with apex of femur and base of tibia dark brown, interspersed pale and dark brown setae present; tibial spurs amber. Basitarsus pale, remaining tarsomeres pale brown, setae brown; distal margins on plantar surface of the first four tarsomeres with dark brown setae; pretarsal claws amber. Wings. Forewing mostly hyaline, dark brown on area surrounding crossveins, M stem and fork, RP stem and stem of its branches, and intermittently on apical twigging near posterior and apical wing margin; membrane on apex of intracubital area strongly marked; major veins and subcostal veinlets alternating pale and dark brown, crossveins dark brown; pterostigma brown with medial pale area; wing margin alternating pale and brown. Hind wing mostly hyaline, with intermittent amber infuscation on the apical twigging of major veins near the posterior and distal margins; pterostigma dark brown with preapical pale area; C, Sc, and R alternating pale and brown, the remainder longitudinal veins, mostly brown; crossveins dark brown; wing margin alternating brown and pale. Abdomen. Tergites mostly brown, pale near the lateral margins, setae mostly pale brown. Some specimens with tergites completely brown; pleural membrane mostly brown, with pale, marginal areas. Sternites pale with lateral, subtrapezoidal brown marks.

Figure 36. 

Plega drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 36). Head. Diamond-shaped in frontal view, rugose, vertexal region raised above compound eyes, with lateral rows of reclined setae; area adjacent to coronal suture and occipital ridge, glabrous, with muscle insertion mark; coronal suture distinct; area adjacent to frontal sutures sunken, ​​supra-antennal region raised, both areas covered with reclined setae; paraocular area concave. Antenna filiform, short, scape 1.5× as long as wide, slightly distally expanded, with short setae, a few longer and thicker on ventral surface; pedicel as long as wide; flagellum slightly dorsoventrally flattened, with 40 or 41 flagellomeres, which are as wide as long; all articles with medial ring of short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus rugose, with fine and short setae. Labrum pentagonal with fine and long setae; maxillary palpus with first palpomere as long as wide, second 1.5× as long as wide, third palpomere 3× as long as wide, fourth palpomeres two times as long as wide, fifth palpomere slightly 1.5× as long as third, with short and thin setae; mentum with thin, long setae. Labial palpus with first palpomere 1.5× as long as wide, second palpomere 4.5 times as long as wide, third palpomere slightly longer than second, palpimacula narrowly ovoid. Thorax. Pronotum slightly longer than wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae, remaining surface with fine, short setae; postfurcasternum ventrally fused, collar-like. Mesonotum 2× as wide as long, with scattered, thick, pedicellate setae and some long, thin setae on medial area. Metanotum ~ 2× as long as wide, mostly glabrous with few fine setae on scutellum. Pteropleura with short, thin setae; a few thick setae on mesanepisternum. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with pedicellate, fine or thick setae of different lengths; trochanter subconical, with thin and short setae, except dorsally with some thick and longer setae, anterior surface with protuberant area covered with pedicellate, thick setae. Femur robust, covered with abundant, fine, short setae which arise from protuberant bases on both surfaces. Closing surface with posteroventral row of processes composed two medially located primary processes and a sub-basal secondary process; between these processes a single tertiary process, and abundant tubercle-shaped processes and stinger-shaped setae are preset; distal portion of posteroventral row raised, composed of tubercle-shaped specializations of different sizes and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½ and distal region; it is composed of tubercle-shaped specializations, stinger-shaped setae, and a basal primary, curved process; distal portion composed tubercle-shaped specializations; adjacent row of thickened setae with globular base present on distal ½. Tibia almost as long as the femur, curved, setose, ventrally keeled with prostrate setae row; a patch of clavate setae is present apically on the anterior surface. Basitarsus with lanceolate process reaching the middle of fourth tarsomere; clavate setae are present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 7× as long as wide; third tarsomere 1.5× as long as wide, fourth tarsomere two times as long as wide. Mid- and hind leg. Coxae with short, thin setae; trochanter on both legs with fine and short setae; femora and tibiae with fine, short setae; tibial spurs short; hind leg longer than midleg, tibia, 1.5× as long as femur; tarsi with fine and short setae, except on distal margin of plantar surface with lateral groups of 5–7 thickened setae; basitarsus 3.5× as long as wide, second tarsomere 1.5× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere two times as long as wide. Wings. Forewing narrowly oval, trichosors present along margin except on proximal ½ of wing; venation setose; costal space proximally expanded, humeral vein branched, 12–14 subcostal veinlets; pterostigma elongated, curved, with incomplete, mostly simple veinlets; subcostal space with single crossvein, medially located; Sc vein abruptly posteriorly curved at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 curved with three or four RP branches arising from it; three veins arising from rarp1; M vein fused basally with RA; RP base widely separated from point of divergence of M from R; M forked slightly before RP base, 1 r-m connecting RP base and MP stem, forming a trapezoidal cell; five or six gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked at level of separation of M and R; A1 apically forked, ending on posterior margin at CuP fork level, A2 forked at CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with 4–6 veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriorly at proximal margin of pterostigma to merge with RA; pterostigma elongated, curved, narrow, composed of incomplete, mostly simple veinlets; radial space with single crossvein, oblique; three or four veins arising from rarp1, 1–3 from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent; CuP not touching A1, strongly anteriorly bent at distal 1/3, with three branches; two crossveins on cubitoanal space; A1 simple ending on wing margin slightly beyond 1m-cu stem level; A2 simple, short and curved. Abdomen. Cylindrical to medially expanded, setae on tergites gradually longer and more abundant towards terminal segments; tergites III-VII with elongated anterolateral scars. Sternites with abundant, thin, short setae; sternites III-VII with lateral scars near the posterior margin.

Male genitalia (Fig. 37A–E). Tergite IX medially narrower than laterally, ventrally bluntly quadrangular. Sternum VIII rectangular; sternite IX U-shaped in ventral view, covered with abundant fine and long setae, posteromedially produced into a blunt lobe, which is equipped dorsal canal; in lateral view short and blunt, the apex reaching posterior margin of ectoproct. Gonocoxites IX thickened, short, sinuous; base spatulate, connected to gonocoxites XI with a membrane; the rest of the structure strongly laterally arched, with three digitiform, apical processes, subequal in shape and size, two of which are dorsally situated; the surface of the digitiform processes is set with, minute, scattered spinules. Ectoproct ovoid, covered with abundant, thin setae; ventral surface concave, forming a broad, rounded lobe, entire surface covered with microtrichia; ventromedially the lobe is continued by a curved, sclerotized canal which led to concave plate. Gonocoxites X forming a short, thickened, ventrally canaliculate sclerite, whose anterior apex is expanded; posterior apex with dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X with a membrane; gonostyli X with thickened, obtuse base, with two lateral processes, the rest of the structure, ribbon-shaped, ventrally curved, and anteriorly recurved, forming a loop before protruding from the abdomen. Gonapophyses X as two short, thin, rod-shaped, straight sclerites arranged in a V-shaped structure; the gonapophyses are joined by a membrane which is medially sclerotized and covers the gonostyli X base. Gonocoxites XI U-shaped, medial lobe complex and elaborated, with two differentiated parts: a dorsal arch and a posteroventral convex area, which is covered with microspinules and medially incised; between these parts a narrow, arched, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI short, sinuous with anterior apex strongly ventrally incurved.

Figure 37. 

Plega drepanicoides Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca.

Female genitalia (Fig. 37F–H). Sternum VII as two medially joined, trapezoidal plates (gonocoxites VII), the fusion line discernible, area adjacent to posterior margin with abundant thick and long setae. Tergite VIII slightly narrower medially than laterally, encircling the spiracle of the segment, lateral margin bluntly quadrangular. Gonocoxites VIII as a trapezoidal concave plate. Gonapophyses VIII as two medially joined trapezoidal plates which form a keel; posteromedial margin of this keel bilobed, with medial U-shaped concavity; median par of gonapophyses VIII connected to lateral rounded plates, which are folded beneath tergite IX + ectoproct. Tergite IX + ectoproct narrowly ovoid. Gonocoxites IX elongated, sinuous, and narrow, as long as the last four abdominal segments combined. Bursa copulatrix long, siphon-shaped, semi-membranous, striated. Spermatheca relatively simple; proximal section, spiral-shaped, thin, with intermittent swellings; medial section short, as wide as proximal section, without swellings; distal section slightly expanded, with short and blunt diverticulum; fertilization canal duct short, triangular, concave, with distal, thin portion; fertilization canal J-shaped, covered with microfilaments.

Distribution

Mexico (Chihuahua, Jalisco).

Remarks

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, 1982

Figs 38, 39

Plega duckei Penny, 1982a: 424. Holotype: male, Brazil, Amazonas (INPA), photographs examined.

Material examined

Holotype. Brazil • 1 ♂; Amazonas, Reserva Ducke; 11 Oct. 1977; J.R. Arias leg.; INPA. Paratypes. Brazil • 1 ♂; Amazonas, Manaus, Parque das Laranjeiras; 22 Jan. 1981; J.R. Arias leg.; “Paratype, Plega duckei Penny”; USNMENT01541899; USNM.

Other material

Brazil • 1 ♀; Amazonas, Manaus; 22 Oct. 1991; G.A.R. Melo leg.; DZUP. • 1 ♀; same data as for preceding; 17 Aug. 1991; DZUP.

Diagnosis

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.

Description

Measurements. Male (n = 1). Forewing length: 8.5 mm; Hind wing length: 6.2 mm.

Coloration (Fig. 38). Head. Vertexal yellow, with broad medial brown area, extended from occiput to supraantennal region, with pale brown setae; area adjacent to coronal suture paler; supra-antennal area darker with medial yellowish area; occiput and postgena pale brown. Antenna brown, with dark brown setae. Frons dark brown, clypeus brown, labrum pale brown; mandible pale brown, with ventral corner dark brown, amber at apex; maxilla with cardo and stipes yellow, galea brown, palpus dark brown; labium with postmentum yellow, pale brown ligula, palpus dark brown. Thorax. Pronotum yellow with brown marks on anterior ½ and posteromedian region; episternum yellow; postfurcasternum pale. Mesonotum with brown marks on sclerites on scutum, pale on area adjacent to sutures; scutellum pale with brown mark on the center; setation mostly pale brown. Metanotum with lateral brown areas and pale medial area; pre-episternum brown; pteropleura brown with small pale areas, setation mostly pale brown. Foreleg. Coxa yellowish with brownish apex; trochanter brownish, setation pale brown. Femur yellowish with pale brown areas; setae mostly pale brown. Tibia dashed with yellowish and pale brown areas, setae pale brown. Basitarsus with yellowish proximal ½ and pale brown lanceolate process; remaining tarsomeres pale brown. Mid- and hind leg. Coxae brown, with pale brown setae; trochanter of both legs yellowish. Femur and tibia of mid-leg alternating yellowing and pale brown, with pale brown setae present. Hind leg with femur yellowish with pale brown apex, setae mostly yellow, tibia alternating pale brown and yellowish, setation mostly pale brown. Tarsi pale brown; first four tarsomeres with brown setae, laterally on distal margin of plantar surface; remainder surface with pale brown setae; pretarsal claws pale brown. Wings. Forewing mostly hyaline; membrane surrounding crossveins on subcostal and radial spaces, RP stem, first two crossveins of mediocubital space and first branch of CuA amber; with intermittent amber areas between apical twigging on distal ½ of wing near posterior and apical wing margins. Pterostigma brown with yellow medial area. Major veins alternating yellowish and brown, CuP and anal veins mostly yellow; subcostal veinlets and crossveins brown; wing margin alternating pale yellowish and brown. Hind wing hyaline, amber on area adjacent to stem first branch of CuA; pterostigma brown with yellowish preapical area; longitudinal veins alternating pale yellowish and brown; subcostal veinlets pale brown; crossveins brown, 1r-m bicolor; wing margin alternating pale yellowish and brown. Abdomen. Cleared.

Figure 38. 

Plega duckei Penny, 1982 A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.

Morphology (Fig. 38). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct; medial area of ​​supra-antennal region slightly raised, with fine, reclined setae; paraocular area concave. Antenna moniliform, long; scape and pedicel as long as wide, both with fine, short setae; flagellum not dorsoventrally flattened, with 44–52 flagellomeres, subconical, progressively narrowed towards the apex, those of proximal ¼ of the flagellum widened; all articles with medial ring of fine, setae. Compound eye hemispherical, as wide as ¾ of the interocular distance at torulus level. Frons rectangular, with fine and short setae, clypeus trapezoidal, narrow with long, thin setae. Labrum pentagonal with thin, short setae; maxillary palpus with first palpomere as long as wide, second 1.2× as long as wide, third palpomere 3× as long as wide, fourth palpomere 2.5× as long as wide, fifth palpomere as long as third, all with short and thin setae; mentum with long, thin setae; labial palpus with first palpomere 2× as long as wide, second palpomere 3.2× as long as wide, third palpomere as long as second, palpimacula grooved. Thorax. Pronotum slightly longer than wide, with slightly raised anterior margin, medial and posterior regions; these areas covered with elongated, thickened setae; remaining surface with fine, short setae. Mesonotum wider than long, with abundant, thick, setae on medial area. Metanotum ~ 2× as wide as long, scutellum with few, fine setae. Pteropleura with short, thin setae. Foreleg. Coxa slightly shorter than femur, cylindrical, anterior and posterior surfaces abundant, thin setae; trochanter narrow, with thin and short setae, except on dorsal surface with some pedicellate, long setae near distal margin; anterior surface without protuberant area covered with pedicellate, long setae. Femur long and narrow, covered with abundant, fine, long setae; closing surface with posteroventral row of processes composed two medially located, primary processes; proximal portion of the row with sub-basal tertiary process; the rest of the row with abundant tubercle-shaped processes, and stinger-shaped setae; distal portion not raised composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal 2/5. Anteroventral row of processes reduced to proximal ½; it is composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of a few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ½. Tibia almost as long as femur, curved, with thin, short setae; ventral surface keeled with prostrate setae; a patch of clavate setae apically on anterior surface is present. Basitarsus with lanceolate process reaching the base of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 7× as long as wide; third tarsomere 1.5× as long as wide, fourth tarsomere 2× as long as wide. Mid- and hind leg. Coxae and trochanter with thin setae, shorter on trochanter; femora with interspersed fine setae of different lengths; hind leg longer than midleg, tibia 1.5× as long as femur, slightly expanded and flattened; tibial spurs short; tarsi with fine and short setae, except on distal margin of plantar surface with lateral groups of 1–3 thickened setae; on both legs, basitarsus 4× as long as wide, second tarsomere 1.5× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere 2× as long as wide. Wings. Forewing oval, trichosors present along margin except on wing base; venation setose; costal space proximally slightly expanded, humeral vein branched, 11 or 12 subcostal veinlets; pterostigma elongated, narrow, gently curved, with incomplete veinlets; subcostal space with single crossvein, medially located; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 gently curved with one or two RP branches; three or four veins arising from rarp1; M vein basally fused with RA; RP base close to divergence of M and R; M forked opposite to RP origin, 1 r-m connecting RP base and M fork, forming a trapezoidal cell; three or four gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked opposite to the level of separation of M and R; A1 simple, ending on posterior margin at level of CuP fork, A2 forked opposite to CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with five veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriad at proximal margin of pterostigma to merge RA; pterostigma elongated, narrow, gently curved, composed of entire veinlets; radial space with single crossvein, oblique; three veins arising from rarp1, one or no veins arising from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein present; CuP not touching A1, anteriorly bent at distal ½, pectinate; two crossveins on cubitoanal space; A1 simple, ending on wing margin at 1r-m stem level; A2 simple, short, and curved. Abdomen. Cylindrical, setae on tergites, scattered, thin, and short; tergites subquadrate. Sternites rectangular, with abundant fine and short setae, longer near the posterior margin of each sternite.

Male genitalia (Fig. 39). Tergite IX medially narrower than laterally; lateral margin rounded. Sternum VIII rectangular; sternite IX U-shaped in ventral view, with short and thin setae; posterior margin with medial, small, blunt lobe which is dorsally canaliculated; in lateral view bluntly trapezoidal, apex reaching posterior margin of ectoproct. Gonocoxites IX short, thin, straight; base flattened, spatulated; apex branched, with three or four short processes. Ectoproct ovoid with short, thin setae, ventrally with broad, rounded, sclerotized lobe, which is continuous with ventromedial, curved sulcus. Gonocoxites X forming a short, ventrally canaliculated sclerite, whose anterior apex is expanded; posterior apex has dorsal processes connected to gonostyli X and ventrolateral processes connected to gonapophyses X with a membrane; gonostyli X with thickened, straight base, equipped two lateral processes, the rest of the structure ribbon-shaped, short, recurved at middle before protruding from abdomen. Gonapophyses X short, thin, straight, with anterior apex slightly expanded, posterior apex dorsally bent; gonapophyses subparallel, joined by a membrane covering the gonostyli X base. Gonocoxites XI U-shaped, medial lobe complex and elaborated, with two differentiated parts: dorsal part as a thin arch; ventral part forming a concave covering; between these parts a broad, circular, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI short, thin, with anterior gently curved. Hypandrium internum tiara-shaped.

Figure 39. 

Plega duckei Penny, 1982 A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral.

Distribution

Brazil (Amazonas).

Remarks

This species was described from Amazonas, Brazil and is known solely from its type locality and adjacent areas (Penny 1982a). Plega duckei was recovered within the clade of South American species as sister of P. paraensis. This is an anomalous species having enlarged compound eyes, discoidal basal, antennal flagellomeres, sulcate palpimacula, and narrow and elongate forefemur. The male genitalia are markedly distinct from other species of the genus with simple gonocoxites XI and short and recurved gonostyli X. The female genitalia of this species remain unknown.

Plega flammata Ardila-Camacho & Contreras-Ramos, sp. nov.

Figs 40, 41

Type locality

Mexico, Baja California Sur: San Venancio, 08 Oct. 1941; Rosa & Bohart leg.

Material examined

Holotype male, pinned. Original label: “Mexico, Baja California Sur, San Venancio, 08 Oct. 1941, Rosa & Bohart leg.”, CAS. Paratypes. Mexico • 1 ♀; Baja California Sur, 9 mi. N. Cabo San Lucas; 09 Sep. 1988; E.G. Riley leg.; blacklight; TAMU-ENTOX0278312; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTOX0278371; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTOX0278336; TAMUIC. • 1 ♂; 3 mi. N. San Antonio; 9–10 Oct. 1968; El. 1200’; E.L. Sleeper & F.J. Moore leg.; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; Sa. Victoria, Casas Viejas; El. 800’; 27–28 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota det. N. Penny, 1998; CAS. • 1 ♂; La Paz, Todos Santos, Km 5 a la Burrera; 23°25'48"N, 110°10'31"W; 145 m; 30 Sep. 2010; C. Mayorga & L. Cervantes leg.; matorral xerófilo, nocturnal collection; CNIN. • 1 ♂; Isla Cerralvo, punta NW; 18 Aug. 1986; F. Arias leg.; CNIN.

Other material

Mexico – Baja California • 1 ♂; San Lorenzo [Island]; N. Banks leg.; MCZ-ENT00681782; MCZ. – Baja California Sur • 1 ♂; Sa. Victoria, Casas Viejas; El. 800’; 27–28 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota det. N. Penny, 1998; CAS. • 2 ♂; 1 mi. SW. Sta. Catarina; 29–30 Jun. 1967; E.L. Sleeper & E.M. Fisher leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; 2.6 miles E. of San Antonio; 30 Jul. 1971, H.G. Real & R.E. Main leg.; U.V. light; CAS. • 1 ♂; 3 mi. N. San Antonio; 9–10 Oct. 1968; El. 1200’; E.L. Sleeper & F.J. Moore leg.; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; Boca de la Sierra; 27–28 Jun. 1967; E.L. Sleeper & E.M. Fisher leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 2 ♂; 4 Km E. El Triunfo (Km 160); 560 m; 05 Sep. 1977; E. Fisher & R. Westcott leg.; CAS. • 1 ♂; 1 mi. East of Migriño; 19 Jul. 1971; H.G. Real & R.E. Main leg.; U.V. light; CAS. •1 ♂; 1 mi. East of Migriño; 17 Jul. 1971; H.G. Real leg.; at U.V. light; CAS. • 1 ♂; 6 mi. E. San José del Cabo; El. 200’; 26–27 Oct. 1968; E. L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 2 ♂; Boca de la Sierra; 27–28 Jun. 1967; E.L. Sleeper & E.M. Fisher leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 2 ♂ 2 ♀; 9 mi SW Hwy 256 on rd to San Isidro; 24°0'55"N, 110°8'0"W; 700 m; 14–15 Jul. 1999; R. Aalbu, K. Brown, I. Stahl & F. Piñero leg.; Blacklite; Plega dactylota, det. N. Penny, 1999; #D00750-1502; CAS. • 1 ♀; San Isidro; 24°00.24'N, 110°09.32'W; 533 m; 15 Aug. 2004; D. Robacker, D. Thomas & J. Burne leg.; TAMU-ENTOX0755721; TAMUIC. • 1 ♂ 1 ♀; 8 mi. SE La Paz; El. 1000’; 13 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; 22 mi. W. La Paz; El. 1000’; 07 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; 9 mi. N San Lucas; El. 1000’; 24–25 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂; 6 mi. NE San Lucas; El. 100’; 25 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♂ 1 ♀; Baja California Sur, 6 mi. E. San José del Cabo; El. 200’; 26–27 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. • 1 ♀; 12 mi. SW. S. José del Cabo; 03 Jul. 1963; E.M. Fisher leg.; Plega dactylota, det. N. Penny, 1998; CAS. • 4 ♂ 16 ♀; Los Cabos, Sierra de la Laguna, Rancho Ecológico Sol de Mayo, Cabañas; 23°29.899'N, 109°47.395'W; 282 m; 15 Aug. 2021; Contreras, Cancino, Luna, Martins & Marquez leg.; light trap; CNIN. •14 ♂; same data as for preceding; 14 Aug. 2021; CNIN. • 10 ♂ 11 ♀; same data as for preceding; Cabañas; 14 Aug. 2021; trampa McPhail/Melaza; CNIN. • 31 ♂ 11 ♀; same data as for preceding; cabaña hacia la cascada; 23°29.890'N, 108°47.415'W; 238 m; 13 Aug. 2021; CNIN. • 4 ♂; same data as for preceding; Vereda arriba cascada en cañón de la zorra; 23°29.910'N, 109°47.720'W; 271 m; 14 Aug. 2021; Luna, Contreras, Barba & Ramírez leg.; light trap; CNIN. • 1 ♂ 3 ♀; same data as for preceding; Vereda arriba cascada en cañón de la zorra; 23°29.910'N, 109°47.720'W; 271 m; 15 Aug. 2021; light trap; CNIN. • 1 ♂ 1 ♀; Valle Perdido, 2 mi N Reserva gate; 23°21'49"N, 110°1'12"W; 500 m; 16–17 Jul. 1999; R. Aalbu, K. Brown, I. Stahl & F. Piñero leg.; Plega dactylota, det. N. Penny, 1999; D00750-1502; CAS. 1 ♀; Sa. Victoria, El Chorro; El. 1000’; 29–30 Oct. 1968; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega dactylota, det. N. Penny, 1998; CAS. 1 ♀; Todos Santos; 18 Oct. 1941; Ross & Bohart leg.; CAS. • 1 ♀; 13 mi E. hwy 19, E. of Todos Santos; 17–18 Sep. 1988; E. Riley; TAMU-ENTOX0278512; TAMUIC. • 1 ♂; La Paz, Todos Santos, Km 5 a la Burrera; 23°25'48"N, 110°10'31"W; 145 m; 30 Sep. 2010; C. Mayorga & L. Cervantes leg.; matorral xerófilo, nocturnal collection; CNIN. • 6 ♂ 1 ♀; La Paz, 7 mi. SW; 06 Aug. 1966; E. & J. Linsley, J. Chemsak, P.D. Hurd leg.; at light; CSCA. • 1 ♀; Km 119 La Paz-San José del Cabo; 13 Sep. 1977; S. Lorna & G. Gutz leg.; CNIN. • 1 ♂; La Paz-San Antonio, “Microondas”; 23°47'24"N, 110°04'35"W; 677 m; 28 Sep. 2010; C. Mayorga & L. Cervantes leg.; matorral xerófilo, nocturnal collection; CNIN. • 2 ♀; 3 mi. N. San Antonio; 9–10 Oct. 1968; El. 1200’; E.L. Sleeper & F.J. Moore leg.; collected at blacklite; Plega fratercula, det. N. Penny, 1998; CAS. • 1 ♀; 3 Km E. La Burrera; 515 m; 02 Sep. 1977; E. Fisher & R. Westcott leg.; CAS. • 1 ♂; 8.4 mi. W. on Ramal a Los Naranjos; 13 Sep. 1988; E.G. Riley leg.; TAMU-ENTOX0278442; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTOX0278450; TAMUIC. • 1 ♀; 8.4 mi W on Ramal a Los Naranjos; 13 Sep. 1988; A.J. Gilbert leg.; CSCA. • 1 ♂; 9.4 mi. W. Hwy 1 on Ramal a San Felipe; 10–11 Sep. 1988; E. Riley; TAMU-ENTOX0278355; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTOX0278379; TAMUIC. • 1 ♂; same data as for preceding; TAMU-ENTOX0278329; TAMUIC. • 1 ♀; same data as for preceding; TAMU-ENTOX0278349; TAMUIC. • 4 ♀; Isla Cerralvo, punta NW; 18 Aug. 1986; F. Arias leg.; CNIN. • 1 ♀; same data as for preceding; Isla Cerralvo; 04 Aug. 1986; CNIN. • 1 ♀; same data as for preceding; L. Cervantes leg.; CNIN. • 1 ♂ 2 ♀; same data as for preceding; 04 Aug. 1986; L. Cervantes leg.; CNIN. • 1 ♀; Isla San José, punta NW; 17 Aug. 1986; F. Arias; CNIN.

Diagnosis

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.

Etymology

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.

Description

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. 40). Head. Vertexal region pale, with lateral dark brown markings extending from occiput to toruli forming a V-shaped pattern, mainly with pale brown setae; area adjacent to occipital ridge dark brown; paraocular area pale with brown areas; supra-antennal area dark brown, with pale brown setae; occiput pale with brown spot, postgena pale sometimes with brown spot. Antennal scape pale, with brown dorsal longitudinal band, entire surface with pale brown setae, and some dark brown setae; pedicel brown, flagellum pale brown with brown setae. Frons either completely brown with or brown with inverted, triangular, pale, medial mark. Clypeus either pale brown or pale with pale brown lateral areas; labrum pale brown; mandible pale with dark brown corners, dark amber at apex; maxilla pale with pale brown areas, palpus brown; labium pale with pale brown areas, setae pale brown; labial palpus brown, pale at junctions, palpimacula pale brown. Thorax. Pronotum pale with irregular brown markings, anterolateral corners with darker spot; episternum brown, sometimes with pale medial area, postfurcasternum pale. Mesonotum with scutum mainly dark brown, except pale on area surrounding sutures, scutellum pale with median and posterolateral dark brown areas, setation mostly dark brown; metanotum mostly dark brown; pre-episternum pale to dark brown; pteropleura mostly dark brown with pale on margins of sclerites, setation pale brown. Foreleg. Coxa mostly pale, with brown spots on base and apex, interspersed pale and dark brown setae present; trochanter pale, setation pale brown, except dorsally with some dark brown setae. Femur posterior surface pale with three weakly to well defined brown spots on medial area, apex brown; anterior surface dark brown, except at base, and area surrounding the primary process. Tibia dashed with pale and dark brown. Basitarsus pale brown proximally, changing to amber towards the apex, clavate setae pale brown; remaining tarsomeres pale brown. Mid- and hind leg. Coxae pale, sometimes with brown areas, setae pale brown; trochanter pale. Femur and tibia of mid-leg dashed with pale and brown, on hind leg with brown at apex of femur and base and apex of tibia. Tarsi pale to pale brown; distal margins on ventral surface with dark brown setae; pretarsal claws brown. Wings. Forewing mostly hyaline; membrane surrounding crossveins, first branch of CuA and apex of CuP amber; posterior and apical margins on distal ½ of wing with intermittent, amber areas between apical branches of longitudinal veins; pterostigma brown with pale medial area; major veins, subcostal veinlets, and wing margin alternating pale and brown. Hind wing hyaline, amber on area adjacent to first branch of CuA; pterostigma brown with pale preapical area; C, subcostal veinlets and Sc with extensive pale areas, remainder longitudinal veins alternating pale and brown; crossveins brown, except sigmoid 1r-m, bicolor; wing margin alternating pale and brown. Abdomen. Tergites pale brown with pale areas, and lateral darker marks; setation mostly pale. Pleural membrane mostly dark brown. Sternites pale with pale brown lateral areas; setation mostly pale.

Figure 40. 

Plega flammata Ardila-Camacho & Contreras-Ramos, sp. nov. A male habitus, dorsal B wings C head, frontal D pronotum, dorsal E forefemur, anterior surface F same, posterior surface.\

Morphology (Fig. 40). Head. Diamond-shaped in frontal view, rugose; vertexal region raised above compound eyes, with lateral rows of reclined setae; area surrounding coronal suture glabrous, with muscle insertion mark; coronal suture distinct; medial area of ​​supra-antennal region not raised, with fine, reclined setae; paraocular area concave. Antenna submoniliform, short; scape 1.5× as long as wide, slightly distally expanded, with fine, short setae; pedicel as long as wide; flagellum dorsoventrally slightly flattened, with 50–60 flagellomeres, which are 2× as long as wide on proximal ¼, and as long as wide on the rest of flagellum; all articles with medial ring of fine, short setae. Compound eye hemispherical, as wide as ½ of the interocular distance at torulus level. Frons and clypeus narrow, with fine and short setae. Labrum pentagonal with thin, short setae; maxillary palpus with first palpomere as long as wide, second 1.5× as long as wide, third palpomere 3× as long as wide, fourth palpomeres 2.5× as long as wide, fifth as long as third, all with short and thin setae; mentum with long, thin setae; labial palpus with first palpomere 2.5× as long as wide, second palpomere 4× as long as wide, third palpomere slightly longer than second, palpimacula narrowly ovoid. Thorax. Pronotum as long as wide, with raised anterior margin, medial and posterior regions; outgrowths covered with pedicellate, thick setae; remaining surface with fine, short setae. Mesonotum 2× as wide as long, with scattered, thick, pedicellate setae on medial area. Metanotum ~ 2× as long as wide, glabrous. Pteropleura with short, thin setae. Foreleg. Coxa as long as femur, cylindrical, anterior and posterior surfaces with pedicellate fine or thick setae of different sizes; trochanter trapezoidal, with thin and short setae, except on dorsal and anterior surfaces with some thickened setae; anterior surface with protuberant area. Femur robust, covered with abundant, fine, short setae; closing surface with posteroventral row of processes composed two medially located, primary processes, sometimes with tertiary process on the middle; proximal portion of the row with two secondary processes; the rest of the row with abundant tubercle-shaped processes, stinger-shaped setae, and some tertiary processes; distal portion raised composed of tubercle-shaped specializations and stinger-shaped setae; adjacent row of thickened setae with globular base present on distal ¼. Anteroventral row of processes reduced to proximal ½; it is composed of tubercle-shaped specializations and stinger-shaped setae; the basal-most primary, curved process is present; distal portion composed of a few tubercle-shaped processes; adjacent row of thickened setae with globular base present on distal ⅔. Tibia almost as long as femur, curved, with thin, short setae; ventral surface keeled with prostrate setae; a patch of clavate setae apically on anterior surface is present. Basitarsus with lanceolate process reaching the middle of fourth tarsomere; clavate setae present proximally on anterior surface; ventrally with single row of prostrate setae; second tarsomere nearly 6× as long as wide; third tarsomere as long as wide, fourth tarsomere 2× as long as wide. Mid- and hind leg. Coxae and trochanter with short, thin setae; femora and tibiae with interspersed fine, short setae and a few thickened setae; tibial spurs short; hind leg longer than midleg, tibia, 1.5× as long as femur; tarsi with fine and short setae, except on distal margin of plantar surface with lateral groups of 5–7 thickened setae; on both legs, basitarsus 3.5× as long as wide, second tarsomere 1.2× as long as wide; third and fourth tarsomeres as long as wide; fifth tarsomere 2× as long as wide. Wings. Forewing narrowly oval, trichosors present along margin except on wing base; venation setose; costal space proximally expanded, humeral vein branched, 9–15 subcostal veinlets; pterostigma elongated, narrow, straight, with distinct veinlets; subcostal space with single crossvein, medially located; Sc vein abruptly posteriorly bent at proximal pterostigma margin to merge with RA; radial space with two crossveins; rarp2 gently curved with three or four RP branches; three veins arising from rarp1; M vein basally fused with RA; RP base, close to divergence of M and R; M forked nearly opposite to RP origin, 1 r-m connecting RP base and M, forming a trapezoidal cell; 3–5 gradate crossveins present. Cubitus deeply forked; CuP basally angled and approaching A1, distally forked slightly beyond the level of separation of M and R; A1 apically forked, ending on posterior margin at level of CuP fork, A2 forked slightly beyond CuP angle level. Hind wing smaller and narrower than forewing, narrowly oval; costal space narrow and reduced, with 5–7 veinlets; C and Sc fused at ¼ of wing length, Sc vein abruptly curved posteriorly at proximal margin of pterostigma to merge RA; pterostigma elongated, narrow, straight, composed of poorly defined veinlets; radial space with single crossvein, oblique; three or four veins arising from rarp1, one from rarp2. 1r-m sigmoid, connecting the stems of M and RP. Media forked beyond R fork. Cubitus deeply forked, intracubital crossvein subparallel to longitudinal wing axis; CuA sinuous, first branch candelabrum-shaped, spur vein absent or present; CuP not touching A1, strongly anteriorly bent at distal 1/3, pectinate; two crossveins on cubitoanal space; A1 simple ending on wing margin at 1m-cu level; A2 simple, short, and curved. Abdomen. Cylindrical to medially expanded, setae on tergites, scattered, thin, and short, gradually longer and more abundant towards terminal segments; tergites subquadrate, tergites III-V with elongated anterolateral scars. Sternites rectangular, with abundant fine and short setae.

Male genitalia (Fig. 41A–E). Tergite IX medially narrower than laterally; lateral margin quadrangular, posterolateral corner with thickened setae. Sternum VIII quadrangular; sternite IX trapezoidal in ventral view, with rounded posterolateral lobes covered with abundant, long setae; posterior margin with short, medial, blunt lobe, which is dorsally canaliculated; in lateral view blunt, apex not reaching posterior margin of ectoproct. Gonocoxites IX thin, sigmoid, long; base flattened, dorsally bent, connected to gonocoxites XI with a membrane; apex thickened, laterally curved, with 8–13 processes arranged in a bunch. Ectoproct ovoid, covered with abundant, thickened setae on posterior surface, posteroventrally with a convex area covered with thin, pedicellate setae; anteroventrally with a flattened, rounded lobe which is continuous with ventromedial sclerotized, curved sulcus. Gonocoxites X forming a short, thickened, ventrally canaliculate sclerite, with anterior apex expanded, posterior apex with dorsal processes connected to gonostyli X and ventrolateral processes; gonostyli X with thickened and straight base with two lateral processes, the rest of the structure, whip-shaped, ventrally curved, and anteriorly coiled, forming two loops before protruding from abdomen. Gonapophyses X rod-shaped, straight, narrow, arranged in a V-shaped structure; gonapophyses joined by a membrane covering the gonostyli X base; this membrane is medially sclerotized and laterally covered with microspinulae. Gonocoxites XI thin, U-shaped, medial lobe complex and elaborated with two differentiated parts: a dorsal, narrow lobe with medial U-shaped concavity; ventral part as a prominent, curved lobe with trapezoidal apex whose surface appears rugose and with microspinulae; between these parts a small, less sclerotized, hyaline area is present; lateral arms of gonocoxites XI straight with anterior apex incurved. Hypandrium internum tiara-shaped.

Figure 41. 

Plega flammata Ardila-Camacho & Contreras-Ramos, sp. nov. A male terminalia, lateral B same, ventral C male genitalia, lateral D same, dorsal E same, ventral F female terminalia, lateral G same, ventral H spermatheca.

Female genitalia (Fig. 41F–H). Sternum VII rectangular with posterior margin concave, setose. Tergite VIII slightly narrower medially than laterally, encircling the spiracle of the segment; lateral margin triangular. Gonocoxites VIII as a narrow sclerite, medially concave. Gonapophyses VIII with medial part boat-shaped, without processes; laterally folded beneath IX + ectoproct, an oval plate is connected to medial part of gonapophyses VIII. Tergite IX-+-ectoproct triangular. Gonocoxites IX elongated, sinuous, and narrow. Bursa copulatrix long, narrow, semi-membranous and striated, except near genital pore, sclerotized. Spermatheca complex and entangled, proximal section, thin, long, forming numerous coils, terminating on a strongly entangled portion; medial section as wide as proximal section, forming a long spiral; distal section as wide as medial section, terminating in an expanded portion, where a blunt diverticulum is present; fertilization canal duct proximally subconical, concave; fertilization canal J-shaped, covered with microfilaments.

Distribution

Mexico (Baja California, Baja California Sur).

Remarks

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

Figs 42, 43

Plega fumosa Linsley & MacSwain, 1955: 16. Holotype: male, Mexico, Michoacán (CAS), specimen examined.

Material examined

Holotype. Mexico • ♂; Michoacán, 11 mi E. Apatzingán; 20 Aug. 1954; E.G. Linsley, J.W. MacSwain, R.F. Smith leg.; Type No. 7202; CAS. Paratypes. Mexico • 1 ♂; Michoacán, 11 mi. E. Apatzingán; 20 Aug. 1954; E.G. Linsley, J.W. MacSwain, R.F. Smith leg.; “Paratype of Plega fumosa Linsley & MacSwain”; FSCA. • 2 ♀; same data as for preceding; FSCA. • 1 ♀; 11 mi. E. Apatzingán; 20 Aug. 1954; E.G. Linsley, J.W. Mac Swain, R.F. Smith leg.; “Paratype Plega fumosa Linsley & MacSwain”; CAS. • 2 ♀; 11 mi. E. Apatzingán; 20 Aug. 1954; E.G. Linsley, J.W. MacSwain, R.F. Smith leg.; MCZ Paratype 29540, “Paratype Plega fumosa Linsley & MacSwain”