With more than 886 species in ∼137 genera, Ectrichodiinae (Hemiptera: Reduviidae) are the largest animal clade of millipede predators. Recent phylogenetic studies have created a framework for our understanding of ectrichodiine evolutionary history, but no fossil species have been described. Ectrichodiella Fracker and Bruner, 1924, belongs to the earliest diverging lineage of Ectrichodiinae, and a better understanding of the morphology and biodiversity of this genus may provide insights into the early evolution of the subfamily. The genus is composed of two described species, Ectrichodiella minima (Valdés, 1910) and Ectrichodiella rafaeli (Gil-Santana and Coletto-Silva, 2005) from Cuba and Brazil, respectively. Here, four new species of Ectrichodiella are described, with three representing extant taxa known only from French Guiana (E. caballina, n. sp., E. nouraguensis, n. sp., and E. obscura, n. sp.), while the fourth is based on a Miocene Dominican amber fossil (Ectrichodiella electrina, n. sp.). Diagnoses, descriptions, habitus and detailed morphological images, an identification key, and a distribution map are provided. A cladistic analysis based on 45 morphological characters that includes the fossil species corroborates the monophyly of Ectrichodiella, but the relationship of the fossil species to the five extant species remains ambiguous. Nevertheless, this phylogenetic placement makes Ectrichodiella electrina, a valuable fossil calibration for future divergence dating analyses, despite its relatively young age.
INTRODUCTION
Ectrichodiinae, the millipede assassin bugs, are a diverse subfamily of Reduviidae (>137 genera, >886 species) of documented or suspected millipede predators (Forthman and Weirauch, 2012; Forthman and Gil-Santana, 2021). Recent phylogenetic analyses have resulted in the first insights into the evolutionary history of the group, including the evolution of aposematic coloration and extreme sexual dimorphism, as well as the colonization of Madagascar (Forthman and Weirauch, 2016, 2017, 2018). These phylogenies were also the basis for a revised classification that treats the former subfamily Tribelocephalinae as a junior synonym of Ectrichodiinae and recognizes five tribes (Forthman and Weirauch, 2017).
The worldwide, mostly circumtropical Ectrichodiini include the bulk of millipede assassin bug species (∼730 spp.) and are often aposematic and glabrous, while the typically brown and setose Tribelocephalini (∼130 spp.) and Abelocephalini (∼20 spp.) are restricted to the Old World. The two remaining tribes form the earliest diverging lineage of Ectrichodiinae and comprise the monogeneric Afrotropical Xenocaucini (8 spp.; Weirauch et al., 2017) and the Neotropical Tribelocodiini (3 spp.; Gil-Santana et al., 2013; Weirauch, 2010). Tribelocodiini currently consist of the monotypic genus Tribelocodia Weirauch, 2010, from French Guiana and Ectrichodiella Fracker and Bruner, 1924, with Ectrichodiella minima (Valdés, 1910) and Ectrichodiella rafaeli (Gil-Santana and Coletto-Silva, 2005) from Cuba and Brazil, respectively (Gil-Santana et al., 2013). The Tribelocodiini are set apart from other Ectrichodiinae by a unique combination of morphological features including the fore- and midtibiae lacking a fossula spongiosa and features of the scutellum, among several other characters (Forthman and Weirauch, 2017). Initially described in the Tribelocephalinae and supported as sister taxon to Xenocaucus China and Usinger, 1949, in a morphology-based phylogenetic analysis (Weirauch, 2010), Tribelocodia was recovered as the sister taxon of Ectrichodiella in the combined morphological and molecular analysis by Forthman and Weirauch (2017). Tribelocodia differs from Ectrichodiella by the 1-segmented tarsus that is fused with the tibia (2-segmented and not fused in Ectrichodiella), absence of dorsal abdominal gland I (present in Ectrichodiella), and a bifid scutellar apex (median scutellar apex and paired midlateral processes in Ectrichodiella) (Forthman and Weirauch, 2017). According to Forthman and Weirauch (2017), the monophyly of Ectrichodiella is supported by several morphological synapomorphies, including the antennal insertion protected laterally by a small sclerite, the toothlike subapical spine on the scape, and paired ventro-apical projections on all femora.
Despite comprising only two described species, the taxonomic history of Ectrichodiella is convoluted. The genus was described by Fracker and Bruner (1924) to accommodate Ectrichodiella cubensis Fracker and Bruner, 1924. This species was later recognized as a junior synonym of Ectoiocchoda minima Valdés, 1910, a misspelling of the manuscript name Ectrichodia minima Uhler (Bruner and Barber, 1937). Ectrichodiella rafaeli was described in a new genus of Reduviinae, Berengeria Gil-Santana and Coletto-Silva, 2005 (Gil-Santana and Coletto-Silva, 2005), and later synonymized with Ectrichodiella (Gil-Santana et al., 2013). Gil-Santana et al. (2013) also provided documentation of type specimens of E. minima and E. cubensis and a discussion of the taxonomic history, morphology, and evidence for its placement in Ectrichodiinae.
While our understanding of the timing of the early evolution of Ectrichodiinae is limited, fossil-calibrated divergence dating analyses by Hwang and Weirauch (2012) and Forthman and Weirauch (2016) suggest that Ectrichodiinae diverged from other assassin bugs in the Late Cretaceous (∼74 mya; HPD ∼85–65 mya) and Tribelocephalini from Ectrichodiini in the early Paleogene (∼62 mya; HPD ∼75–50 mya). While Xenocaucini and Tribelocodiini were not included in these analyses, their phylogenetic position as early-diverging lineages of Ectrichodiinae makes it likely that they originated close to the K-Pg boundary (Forthman and Weirauch, 2017). Biogeographic analyses of Madagascan Ectrichodiini show that this fauna is young and the result of dispersal-driven colonization during the Paleogene (Forthman and Weirauch, 2016). In contrast, given the potential age of Tribelocodiini and the distribution of extant taxa in the Guiana Shield, Amazonia, and the Greater Antilles, vicariance via the Greater Antilles and Aves Ridge (GAARlandia) land bridge (35–32 Mya) (Iturralde-Vinent, 2006) may have impacted the biogeographic history of this tribe.
Fossil species provide insights into the evolution of morphological characters and the timing of lineage diversification when analyzed in a phylogenetic framework. Despite this, no work has provided a comprehensive background examination of current knowledge of extinct Reduviidae. While Reduviidae comprise more than 7,000 described extant species, Schuh and Weirauch (2020) recognized only ∼34 fossil species as belonging to this family. Emesinae are best represented (13 spp.) in the fossil record, followed by Holoptilinae (3 spp.), Triatominae (3 spp.), Harpactorinae (2 spp.), Reduviinae (2 spp.), and Centrocnemidinae (2 spp.), while subfamilial placement of the remaining fossil taxa is uncertain. Fossil ectrichodiine taxa have not been documented. Most assassin bug fossils are fairly young (Eocene, Oligocene, and Miocene) and comprise compression fossils from deposits including Florissant, Messel, and Shandong and amber fossils from Baltic and Dominican amber (Schuh and Weirauch, 2020). The recently described Paleotriatoma Poinar, 2019, is the first assassin bug from mid-Cretaceous Burmese amber and oldest uncontroversial fossil Reduviidae (Poinar, 2019). While Dominican amber fossils are young (Iturralde-Vinent, 2017), they are easily placed within a given subfamily and tribe and are often part of extant genera. Dominican amber reduviid fossils comprise species of Emesinae, Harpactorinae (Apiomerini), Holoptilinae, and Triatominae. Certain extant Emesinae (Wygodzinsky, 1966) and Holoptilinae (C.W., personal obs.) are found around the bases of trees and Apiomerini collect resin for maternal care and prey capture (Forero et al., 2011), circumstances that likely facilitate fossilization in amber. Ectrichodiinae are often collected from leaf litter (Forthman et al., 2016) or on vegetation (Haviland, 1931), and while most photographs available on community science portals like iNaturalist.org ( https://www.inaturalist.org) show millipede assassin bugs on artificial surfaces or the ground, some were observed on tree bark. Thus, their absence from the amber fossil record is surprising.
We here describe and illustrate four new species of Ectrichodiella and provide an updated diagnosis and description of the genus. Three of the new species are extant taxa from French Guiana, and the fourth is the first fossil species of Ectrichodiinae and is from Dominican amber. We conduct a morphology-based phylogenetic analysis of Ectrichodiella and outgroups to reconstruct the phylogeny of the genus and determine the phylogenetic position of the fossil species.
MATERIALS AND METHODS
Specimens, Databasing, and Map
Between four and nine male specimens and one or two female specimens were examined for each of the three new extant species and E. rafaeli, and one male specimen was examined for the new fossil species (figs. 1–14). Specimens of extant species were collected via window trap by the Société Entomologique Antilles-Guyane (SEAG). The Dominican amber specimen is deposited at the American Museum of Natural History, New York (AMNH). Holotypes of the three extant species are deposited at the Muséum National d'Histoire Naturelle, Paris (MNMH) and paratypes in the collection of Jean-Michel Berenger (JMB), MNHN, and the Entomology Research Museum at the University of California, Riverside (UCR).. Since E. minima is well described and documented, type specimens were not examined. Examined specimens were given matrix-code labels (unique specimen identifier or USI labels) and specimen data captured using the online Arthropod Easy Capture (AEC) database maintained at the AMNH ( https://research.amnh.org/pbi/locality/). Records are publicly available through the Heteroptera Species Pages ( https://research.amnh.org/pbi/heteropteraspeciespage/). Coordinates were downloaded from the AEC database and mapped using SimpleMappr ( http://www.simplemappr.net/) (fig. 15).
Dissections, Imaging, and Measurements
Morphological characters were examined using a Nikon SMZ1000 dissecting microscope. The pygophore of the four extant species was dissected and cleared in ∼10% warm potassium hydroxide and the aedeagus removed. Genitalic dissections were stored in genitalic vials with glycerol and re-associated with the specimen after examination and documentation. Habitus and detailed morphological photographs (figs. 1–14) were taken using a Leica Z16 APO imaging system with 1.0× and 2.0× objective lenses and the Leica Application Suite v4.3 software and compiled using Zerene Stacker v1.04. Photographs were edited and plates assembled in Adobe Photoshop CS6. Measurements (in mm) were taken from the photographs. Holotype measurements are provided as part of the description. See table 1 for a list of measurements.
FIG. 1.
Habitus of Ectrichodiella spp. in dorsal, lateral, and ventral views. A–C. E. caballina, n. sp.; D–F. E. nouraguensis, n. sp.; G, H. E. electrina, n. sp.; I–K. E. obscura, n. sp.; L–N. E. rafaeli.
FIG. 2.
Head in dorsal (A, C, E) and lateral (B, D, F) views. A, B. E. caballina; C, D. E. electrina; E, F. E. nouraguensis. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: as, antennal shield; gl, gula; L2, labial segment 2 (first visible); L3, labial segment 3 (second visible); L4, labial segment 4 (third visible); oc, ocellus; psp, prosternal stridulatory process; sas, subapical spine on antennal scape.
FIG. 3.
Head in dorsal (A, C) and lateral (B, D) views. A, B. E. obscura; C, D. E. rafaeli. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: as, antennal shield; gl, gula; L2, labial segment 2 (first visible); L3, labial segment 3 (second visible); L4, labial segment 4 (third visible); oc, ocellus; sas, subapical spine on antennal scape.
Terminology
Morphological terminology follows Forthman and Weirauch (2016), with modifications.
Abbreviations: 1A, first anal vein; alp, anterior pronotal lobe processes; as, antennal shield; asp, apical scutellar process; bp, basal plate (of aedeagus); bpe, basal plate extension; ca, carina (median longitudinal of abdomen); Cu, cubitus; DAG1–3, dorsal abdominal gland 1–3; exM, extension of M beyond M+Cu; gl, gula; he, hemispherical elevations anterior of interocular sulcus; L2, labial segment 2 (first visible); L3, labial segment 3 (second visible); L4, labial segment 4 (third visible); lr, lateral ridges of posterior pronotal lobe; M, media; mp, midlateral projections of scutellum; msn, mesonotum; mtn, metanotum; oc, ocellus; pa, paramere; pg, prosternal groove; ple, paramedian lobe-like extension on posterior margin of posterior pronotal lobe; pr, pygophore ridge (median on ventral pygophore surface); psp, prosternal stridulatory process; pt, pterostigma; py, pygophore; S3–S8, (abdominal) sternites 3–8; sas, subapical spine on scape, sc, scutellum; T1–2, synterga 1 and 2; T3–T7, (abdominal) terga 3–7; td, (fore) tibial dilation; ts1, 2, tarsal segment 1, 2; vap, paired ventroapical projections on forefemur.
Species Descriptions and Cladistic Analyses
To assure consistency of species descriptions, descriptive characters and states were organized in Microsoft Excel and then assembled into text sections and edited in Microsoft Word. The morphological matrix (supplementary file S1 available online: https://doi.org/10.5531/sd.sp.69) used for phylogenetic analyses overlaps with characters used for the descriptions. However, unique species-level characters were omitted from the phylogenetic matrix, and additional characters were included that do not vary between species of Ectrichodiella and are aimed at testing the monophyly and relationship of Ectrichodiella. The morphological matrix was assembled in WinClada ver. 1.00.08 (Nixon, 2002) and comprises 45 morphological characters and 11 taxa: two previously described and four new species of Ectrichodiella, Tribelocodia ashei Weirauch, 2010 (Tribelocodiini), Tribelocephala sp. (Tribelocephalini), Abelocephala sp. (Abelocephalini), Xenocaucus ethiopiensis Weirauch et al., 2017 (Xenocaucini) and Gibbosella consimilis Forthman, Chłond, and Weirauch, 2017 (Ectrichodiini). Because of the exceptional preservation of the fossil specimen, 40 of the 45 characters were coded for this taxon.
FIG. 6.
Lateral (A, C, E, G) and ventral (B, D, F, H) views of labium. A, B. E. caballina; C, D. E. nouraguensis; E, F. E. obscura; G, H. E. rafaeli. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: L2, labial segment 2 (first visible); L3, labial segment 3 (second visible); L4, labial segment 4 (third visible); psp, prosternal stridulatory process.
FIG. 7.
Dorsal (A, C, E) and lateral (B, D, F) views of thorax. A, B. E. caballina; C, D. E. electrina; E, F. E. nouraguensis. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: alp, anterior pronotal lobe processes; asp, apical scutellar process; lr, lateral ridges (of posterior pronotal lobe); mp, midlateral projections of scutellum; msn, mesonotum; mtn, metanotum; pg, prosternal groove; ple, paramedian lobe-like extensions on posterior margin of posterior pronotal lobe; sc, scutellum; ts1, ts2, tarsal segments 1 and 2.
FIG. 8.
Dorsal (A, C) and lateral (B, D) views of thorax. A, B. E. obscura; C, D. E. rafaeli. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: alp, anterior pronotal lobe processes; asp, apical scutellar process; lr, lateral ridges (of posterior pronotal lobe); mp, midlateral projections of scutellum; ple, paramedian lobe-like extensions on posterior margin of posterior pronotal lobe; sc, scutellum.
Cladistic analyses were implemented as equal and implied weights. For equal weights (EW) analysis in TNT (Goloboff et al., 2008), we used New Technology Searches, using sectorial Searches, ratchet, drift, and tree fusing with default settings, and finding the minimum-length tree 100 times. For resampling procedures, we employed jackknife with default settings and 1000 replications. A number of characters showed homoplasy in the equal weights analysis, so we downweighted homoplastic characters exploring a range of k-values (20, 12, 9, and 6, with 20 downweighting homoplasy the least and 6 the strongest) in implied weights (IW) analyses (Goloboff, 1993). We used symmetric resampling procedures with default settings and 1000 replications to assess the robustness of these analyses.
PHYLOGENETIC RESULTS AND DISCUSSION
The EW analysis (fig. S2A) resulted in one tree (L = 86; Ci = 67, Ri = 72). Ectrichodiella (98% jackknifing) was highly supported in resampling analyses, but relationships within the genus were less supported. The fossil species Ectrichodiella electrina, n. sp., was recovered with poor support as the sister of Ectrichodiella obscura, n. sp. + (Ectrichodiella caballina, n. sp. + Ectrichodiella nouraguensis, n. sp.) (68%), which together were sister to (E. minima + E. rafaeli) (78%).
FIG. 9.
Legs. A. Foreleg, lateral view; B. Foreleg, trochanter, ventral view; C. Foreleg, paired ventroapical projections, ventral view; D. Foreleg, tarsus and pretarsus, lateral view. A, C. E. caballina; B, D. E. nouraguensis. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: td, (fore)tibial dilation; ts1, ts2, tarsal segments 1 and 2; vap, paired ventroapical projections on forefemur.
FIG. 10.
Hemelytron. A. E. caballina; B. E. nouraguensis; C, D. E. electrina; E. E. obscura; F. E. rafaeli. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: 1A, first anal vein; Cu, cubitus; exM, extension of M beyond M+Cu; M, media; pt, pterostigma; Sc, subcosta.
The four IW analyses resulted in one optimal tree, with identical relationships among them after jackknifing, suggesting that homoplasy has little impact on this dataset. Ectrichodiella was monophyletic, E. minima and E. rafaeli were sister taxa, and the fossil species Ectrichodiella electrina, was sister to them (fig. 16; k = 20). Ectrichodiella obscura, was sister to (E. caballina + E. nouraguensis). Ectrichodiella electrina, n. sp. + (E. minima + E. rafaeli) was sister to E. obscura + (E. caballina + E. nouraguensis). In contrast, both the EW and IW before jackknifing (fig. S2) showed E. electrina, as sister taxon to E. obscura (E. caballina + E. nouraguensis).
FIG. 11.
Ventral (A, C, E) and lateral (B, D, F) views of abdomen. A, B. E. caballina, C, D. E. electrina, E, F. E. nouraguensis. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: ca, carina; DAG1, dorsal abdominal gland 1; DAG2, dorsal abdominal gland 2; DAG3, dorsal abdominal gland 3; py, pygophore; S3–8, sterna 3-8; T1–2, synterga 1 and 2; T3–7, terga 3–7.
FIG. 12.
Ventral (A, C) and lateral (B, D) views of abdomen. A, B. E. obscura, C, D. E. rafaeli. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: py, pygophore; S3–8, sterna 3–8.
Across analyses, relationships among the outgroups differed among analyses, which was driven by the phylogenetic placement of Tribelocodia ashei. The phylogenetic position of the fossil species within Ectrichodiella also remains unresolved, given the conflicting position among analyses and very low branch support. We use the tree with the best fit score (1.307, derived from the k = 20 IW analysis; fig. 16) to document unambiguous character state transitions (homoplasy setting: any extra step) and for the Caracter List and Discussion below.
Character List and Discussion of Character Transitions and Synapomorphies Derived from the Implied Weighting (k = 20) Analysis
Characters and states are documented on figures 1–14. Only unambiguous character state changes are discussed (fig. 16). The most recent common ancestor of Ectrichodiella is abbreviated as MRCA.
0. Postocular area, shape in dorsal view: 0, narrow (fig. 2A); 1, broad. A narrow postocular area independently evolved in E. rafaeli and the outgroups (fig. 3C).
1. Posterolateral gular area, shape in lateral view: 0, flat to slightly swollen, conforming to rounded head shape; 1, moderately swollen; 2, distinctly swollen. A distinctly swollen posterolateral gular area (figs. 2B, F; 3B, D) is a synapomorphy for Ectrichodiella.
2. Labium, relative length of second (L2) and third (L3) segments: 0, L3 about 1/2 as long as L2; 1, L3 about 3/4 as long as L2; 2, about same length. The MRCA was reconstructed as having L3 about 3/4 the length of L2 (fig. 6A), with transitions to half the length within the genus (fig. 6C). We were unable to code this character for E. electrina, where it is obscured.
3. Eye, shape, in dorsal view: 0, with close to 90° angle (fig. 2A, E); 1, with distinctly greater than 90° angle (fig. 3A, C). The eye shape in dorsal view shows homoplastic transitions within Ectrichodiella and the outgroups. The reconstruction of the MRCA is ambiguous.
4. Eye, size, in lateral view relative to ventral head margin: 0, not reaching ventral head margin (fig. 2B); 1, reaching ventral head margin; 2, surpassing ventral head margin. The reconstruction for the MRCA was ambiguous. Only E. minima and E. electrina, have eyes that reach the ventral head margin among species of Ectrichodiella.
5. Eye, shape in lateral view: 0, reniform; 1, oval. The eye was reconstructed as being reniform in the MRCA of Ectrichodiella (fig. 3D), with one transition to oval in E. electrina.
6. Synthlipsis (min. interocular distance), width relative to eye: 0, synthlipsis 2× width of eye; 1, synthlipsis 3× width of eye; 2, synthlipsis 1.5× width of eye; 3, synthlipsis more than 3× width of eye. The synthlipsis being more than 3× as wide as the eye is a synapomorphy of (E. minima + E. rafaeli) (fig. 3C). The synthlipsis is narrower in E. caballina (3× eye width; fig. 2A), and is narrowest E. electrina, E. obscura, and E. nouraguensis (2× eye width; fig. 2E).
7. Ocellar lens: 0, absent; 1, present. The presence of the ocellar lens is plesiomorphic for Ectrichodiella and was retained in all species (fig. 2A, C, E; 3A, C).
8. Ocellus, size: 0, small (distance between ocellus and eye margin ∼1.5× ocellar lens diameter); 1, large (distance between ocellus and eye margin ∼1× ocellar lens diameter). Among species of Ectrichodiella, only E. rafaeli shows a relatively small ocellus (fig. 3C). The ocellus is large in the remaining species (fig. 2A, C, E) and is synapomorphic for Ectrichodiella.
9. Ocellus, located on elevation: 0, absent, located on flat cuticle; 1, present. The ocellus in all species of Ectrichodiella except E. rafaeli is located on an elevation (figs. 2B, F; 3D), and this was reconstructed as the state in the MRCA of the genus.
10. Ocellar elevation (if present): 0, single median elevation; 1, paired elevations. The ocelli in E. electrina are located on a single, median elevation (fig. 2C), while those in E. caballina, E. nouraguensis, E. obscura, and E. minima are located on separate, paired elevations (fig. 2A, F).
11. Antennal shield (i.e., antennal insertion shielded laterally by small sclerite): 0, absent; 1, present. The antennal shield (as) is present in all species of Ectrichodiella (figs. 2B, D, F; 3A, C) and a synapomorphy of the genus.
12. Scape of antenna, subapical spine: 0, absent; 1, small; 2, large. A large subapical spine on the scape is a synapomorphy for Ectrichodiella. It is retained in E. electrina, E. caballina, E. nouraguensis, and E. obscura, (fig. 4B, D, F). The spine is small in both E. rafaeli and E. minima (fig. 5D).
13. Flagellomeres, diameter: 0, similar to diameter of pedicel; 1, more slender than diameter of pedicel. Slender flagellomeres are plesiomorphic for Ectrichodiella (figs. 4A, C, E; 5A, C).
14. Anterior and posterior pronotal lobes, relative length: 0, anterior lobe less than 1/2 length of posterior; 1, anterior lobe more than 1/2 length of posterior. The longer anterior pronotal lobe is a synapomorphy for E. electrina, E. rafaeli, and E. minima (fig. 8B). The remaining Ectrichodiella species have a very short anterior lobe (fig. 7A, E) that is here treated as homologous to the condition seen in the sister clade.
15. Anterior and posterior pronotal lobes, relative width: 0, anterior lobe less than 1/2 width of posterior; 1, anterior lobe more than 1/2 width of posterior. The MRCA of Ectrichodiella had a pronotum with the anterior lobe more than 1/2 the width of the posterior lobe, a feature that was retained in the four new species (figs. 7A, C, E; 8A). The anterior lobe being less than 1/2 the width of the posterior lobe is a synapomorphy for (E. minima + E. rafaeli) (fig. 8C).
16. Anterior pronotal lobe, longitudinal depression: 0, absent; 1, present. All species of Ectrichodiella except E. electrina show a longitudinal depression on the anterior lobe. The feature was reconstructed as plesiomorphic for the genus in our analysis.
17. Anterior pronotal lobe, pair of glabrous areas posterior to anterior lobe process: 0, absent; 1, present. The glabrous areas (fig. 7A) are a synapomorphy for Ectrichodiella (state in E. electrina unknown).
18. Anterior pronotal lobe, paired, rounded processes on disc: 0, absent; 1, present. Paired rounded processes on the anterior pronotal lobe disc are a synapomorphy for Ectrichodiella (figs. 7A, C, E; 8A, C).
19. Posterior pronotal lobe, coloration: 0, pale brown; 1, brown. The MRCA of Ectrichodiella was reconstructed with brown posterior pronotal lobe (figs. 7A, C, E; 8A, C); the light brown color in E. minima is autapomorphic.
20. Posterior pronotal lobe, median groove: 0, absent; 1, present. A longitudinal groove is plesiomorphically present in all species of Ectrichodiella (fig. 7A, C, E).
21. Posterior pronotal lobe, lateral ridges (beyond basal half of posterior lobe): 0, absent; 1, present. Lateral ridges (lr) occur in the four new species of Ectrichodiella (fig. 7A, C, E) and may or may not be independently derived.
22. Posterior pronotal lobe, paramedian lobes on posterior margin: 0, shallow; 1, pronounced. Pronounced paramedian lobes are a synapomorphy for Ectrichodiella.
23. Prosternal stridulatory process, length: 0, short, not reaching or surpassing posterior margin of procoxal cavity; 1, long, surpassing posterior margin of procoxal cavity. The process is plesiomorphically long in all species of Ectrichodiella except E. rafaeli for which this feature was examined (unknown in E. minima).
24. Scutellum, apex: 0, drawn into process; 1, blunt. The optimization of this character is ambiguous, with a median process being present in Tribelocephalini, Xenocaucus, and Abelocephala, blunt in Ectrichodiella (fig. 7A, C, E), and absent in Ectrichodiini (though occurring in some species of Ectrichodiini, it does not occur in the species sampled in this study) and Tribelocodia.
25. Scutellum, midlateral projections: 0, absent; 1, paired. All species of Ectrichodiella feature distinct paramedian scutellar projections or processes (mp; figs. 7A–C, E, F; 8A–D). Our analyses found the subapical lateral projections on the scutellum in Tribelocodia to be homologous with the projections in Ectrichodiella.
26. Scutellum, midlateral projections, size: 0, small; 1, large. The four new species share the large size of the midlateral projection (fig. 7A, C, E), here treated as plesiomorphic, because it also occurs in Tribelocodia. The small size of the projections is synapomorphic for E. minima and E. rafaeli species.
27. Scutellum, midlateral projections, orientation: 0, mostly horizontal; 1, strongly vertical. While the projections are plesiomorphically horizontal in the two previously described species of Ectrichodiella and in E. electrina the strong vertical orientation (fig. 7B, F) is a synapomorphy for E. caballina, E. obscura, and E. nouraguensis.
28. Hind trochanter, small tubercles: 0, absent; 1, present. Small tubercles on the hind trochanter are here treated as a synapomorphy for Ectrichodiella (unknown in E. electrina) and Tribelocodia.
29. Forefemur, paired ventroapical projections: 0, absent; 1, present. Paired ventroapical projections on the forefemur are a synapomorphy for Ectrichodiella (vap; unknown in E, electrina; fig. 9A, C).
30. Foretibia, dilation at apex: 0, absent; 1, present. The foretibial apex is dilated (fig. 9A) in all species of Ectrichodiella, except E. electrina. The reconstruction for the MRCA is ambiguous due to the absence of a dilation in Tribelocodia.
31. Fossula spongiosa on foreleg: 0, absent; 1, present. The fossula spongiosa is plesiomorphically absent in all species of Ectrichodiella (fig. 9D).
32. Tarsal segmentation: 0, 2-segmented; 1, 3-segmented; 2, 1-segmented. The 2-segmented tarsus (fig. 9D) is here treated as plesiomorphic for Ectrichodiella (shared with Xenocaucus and Abelocephala).
33. M and Cu (forewing), proximal portion: 0, fused into M+Cu; 1, separate veins. The proximal portions of M and Cu are distinct in the four new species described here, and then fuse in a more distal point (fig. 10). This condition is synapomorphic (but homoplastic), and in contrast to the two previously described species that retain the plesiomorphic condition of a completely fused M+Cu.
34. Pterostigma (inflated area on forewing margin anterior to M): 0, slightly inflated; 1, strongly inflated; 2, not inflated. A strongly inflated pterostigma is synapomorphic (fig. 16) for E. electrina + (E. minima + E. rafaeli) (fig. 10C, D, F).
35. Vestiture of corium: 0, long, simple setae absent; 1, long, simple setae sparse; 2, long, simple setae dense. The long and sparse setation on the forewing is here treated as plesiomorphic for Ectrichodiella (fig. 10).
36. Corium, color: 0, uniformly brown; 1, brown with yellow band proximally. While the corium is plesiomorphically brown in the two previously described species and E. electrina. (fig. 10C), the brown and yellow patterning in E. caballina and E. nouraguensis, is a synapomorphy for two of the new species (fig. 10A, B).
37. Corium, vein color: 0, uniformly yellow; 1, brown and yellow; 2, uniformly brown. The uniform yellow coloration of the veins in E. minima and E. rafaeli is synapomorphic.
38. Lateral margin of laterotergite 1+2: 0, with small spine; 1, with large, curved spine; 2, only slightly expanded; 3, without spine or expansion. None of the outgroups feature a spine or distinct expansion on laterotergite 1+2. The size of the spine varies among species of Ectrichodiella. While the laterotergite is only slightly expanded in the four new species (figs. 11A, C, E; 12A), the spine is large in E. rafaeli (fig. 12C), and small in E. minima.
39. Dorsal abdominal scent gland (DAG) 1 ostioles in adults: 0, absent; 1, present. The DAG 1 ostioles are present (fig. 11C) in species of Ectrichodiella (as are the ostioles of DAG 2 and 3), but it is unclear whether they are synapomorphic or plesiomorphic (absent in Tribelocodia).
40. Connexivum, color pattern: 0, uniformly brown or yellow; 1, with contrasting brown and yellow markings. The contrasting connexival color pattern is a synapomorphy for E. caballina, E. nouraguensis, and E. obscura (fig. 11A, E).
41. Connexivum, color pattern, dark/pale proportion: 0, dark and pale of similar dimension and both square; 1, dark dominant, pale as slender triangle. This feature is a species-diagnostic difference for E. nouraguensis and E. obscura and E. caballina (fig. 1B, E, J).
42. Posterior margin of tergum 7 (male): 0, straight; 1, indented; 2, slightly rounded. The posterior margin of tergum 7 shows some homoplastic transitions among species of Ectrichodiella and can be used to distinguish among the new species and E. rafaeli from French Guiana (fig. 11A, E; 12A).
43. Sternum 8 (male), posterior margin: 0, strongly sinuous; 1, slightly sinuous; 2, straight. This feature shows diagnostic differences between Ectrichodiella species but does not group species together (fig. 13A, H, P, Q, X). It is unknown in E. minima.
44. Pygophore, shape of posterior margin (in ventral view): 0, rounded; 1, straight; 2, slightly rounded. Similar to the preceding character, the posterior pygophore margin differs among species of Ectrichodiella (fig. 13A, I, P, Q, X).
TAXONOMY
Ectrichodiella Fracker and Bruner, 1924
Figures 1–14
Ectrichodiella Fracker and Bruner, 1924: 17: 167. Type species: Ectrichodia [sic] minima Valdés, 1910 (by monotype).
Berengeria Gil-Santana and Coletto-Silva, 2005: 35: 298; junior synonym: Gil-Santana et al., 2013.
Revised diagnosis. Recognized among Ectrichodiinae by the small size, antennal insertion with antennal shield (fig. 2B, F), posterolateral gular area very swollen in lateral view (fig. 2B), anterior pronotal lobe with pair of processes on disc and glabrous areas posterior to processes (fig. 7A, C, E), pronounced paramedian lobe-like extension on posterior pronotal margin (fig. 7A, C, E), scutellum with apex drawn into process (fig. 7A, C, E), midlateral scutellar projections (fig. 7A–C, E, F), forefemur with paired ventroapical projections (fig. 9A, C), absence of fossula spongiosa on fore- and midlegs (fig. 9D), 2-segmented tarsi (fig. 9D), strongly inflated pterostigma in some species (fig. 10C), sparse, long vestiture on corium, and dorsal abdominal scent gland (DAG) ostioles 1–3 present in adults (fig. 11C).
Redescription. Male: Macropterous, small body size (length clypeus to abdominal margin: 3.31–3.74). COLORATION: Body and appendages brown to light brown or yellow; hemelytron and veins brown and/or yellow; connexivum either uniformly colored or with contrasting yellow and brown bands. VESTITURE: Head with light colored setation; antennal vestiture length and hue variable; thorax with light setae, scutellum with long or short, thin, yellow setae; hemelytron with sparse setation. STRUCTURE: HEAD: Antennal insertion protected laterally by antennal shield; interocular sulcus distinct throughout, most pronounced at middle of head, or most pronounced anterior to ocelli; paired hemispherical elevations anterior of interocular sulcus present or absent; synthlipsis ≥2× eye width; posterolateral gular area very swollen in lateral view; eye large (reaching ventral head margin in lateral view) or relatively large (not reaching ventral head margin in lateral view), reniform or oval in lateral view; ocelli size variable, on shared or individual paired elevation(s) or on flat dorsal head surface. Antenna: scape with or without subapical spine, if present, spine large or small; scape and pedicel thickened; flagellomeres more slender than pedicel. Labium: fairly stout, third segment (second visible) ≥0.5× length of second segment (first visible), fourth segment (third visible) reaching prosternum. THORAX: Pronotum bell shaped in dorsal view, with anterior lobe shorter and narrower than posterior lobe, surface either smooth or granulose; anterior pronotal lobe size variable, with anterior margin rounded or straight, with or without median longitudinal depression, with paired, rounded processes on disc and with or without pair of glabrous areas posterior to processes; posterior pronotal lobe with median longitudinal groove, with or without lateral ridges, and with paramedian lobes on posterior margin; scutellum triangular with apex drawn into median, apical process, midlateral scutellar projections small or large, mostly horizontal or strongly vertical. Legs: hind trochanter with small tubercles; forefemur with paired ventroapical projections (fig. 9C); fossula spongiosa on fore- and midlegs absent; foretibia with or without apical dilation, with foretibial comb; tarsi 2-segmented. Hemelytron: membrane with two closed cells (MCu and Cu1A cells) of slightly unequal size (MCu larger), with distal margin of MCu cell curved and distal margin of CuA1 cell fairly straight (fig. 10A, B, D) or slightly curved; proximal portion of M+Cu separated or fused; pterostigma strongly to slightly inflated. ABDOMEN: Lateral margin of laterotergite 1+2 slightly expanded or with small or large, curved spine; posterior margin of tergum 7 straight, indented, or slightly rounded; sterna with median keel (not observed in E. minima), posterior margin of sternum 8 straight to strongly sinuous (fig. 13A, H); ostioles of dorsal abdominal scent glands (DAG) 1–3 present in adults. Genitalia (fig. 13): pygophore rounded (fig. 13P) or more elongate and with sinuous lateral margins (fig. 13A, B, H, I), ventral surface with distinct setation and ridges along midline in some species (fig. 13H), posterior margin of pygophore slightly rounded (fig. 13H) or distinctly rounded (fig. 13O). Paramere slender, of fairly uniform diameter, and gently curved. Aedeagus simple and slender (fig. 13E–G, L–N).
FEMALE: Overall habitus, coloration, and size as male.
FIG. 13.
Male genitalic structures of E. caballina, E. nouraguensis, and E. electrina. A–G. E. caballina; A. Ventral external morphology, B–D. Pygophore ventral, dorsal, lateral views respectively, E–G. Aedeagus dorsal, lateral, ventral views respectively; H–N. E. nouraguensis, H. Ventral external morphology, I–K. Pygophore ventral, dorsal, lateral views respectively, L–N. Aedeagus dorsal, lateral, ventral views respectively; O, P. Ventral external morphology E. electrina Numbers refer to characters and character states used in cladistic analysis. Abbreviations: bp, basal plate; bpe, basal plate extension; ca, carina; pa, paramere; pr, pygophore ridges; S8, sternite 8.
FIG. 14.
Male genitalic structures of E. obscura, and E. rafaeli. A–G. E. obscura; A. Ventral external morphology, B–D. Pygophore ventral, dorsal, lateral views respectively, E–G. Aedeagus dorsal, lateral, ventral views respectively; H–N. E. rafaeli; H. Ventral external morphology, I–K. Pygophore ventral, dorsal, lateral views respectively, L–N. Aedeagus dorsal, lateral, ventral views respectively. Numbers refer to characters and character states used in cladistic analysis. Abbreviations: bp, basal plate; bpe, basal plate extension; pa, paramere; S8, sternite 8.
Key to Species of Ectrichodiella
1. Lateral margin of laterotergite 1+2 with large, curved spine (fig. 12C); ocelli not on elevation (fig. 3D) E. rafaeli
– Lateral margin of laterotergite 1+2 with small spine or only slightly expanded; ocelli on elevation 2
2. Midlateral projection of scutellum small, posterior pronotal lobe without lateral ridges, lateral margin of laterotergite 1+2 with small spine E. minima
– Midlateral projection of scutellum large (fig. 7B, C, F), posterior pronotal lobe with lateral ridges (fig. 7A, C, E), lateral margin of laterotergite 1+2 slightly expanded 3
3. Ocelli on shared elevation (fig. 2C), midlateral scutellar projections mostly horizontal (fig. 7C), pygophore rounded (fig. 13P) E. electrina.
– Ocelli on individual elevations (fig. 2A, E), midlateral scutellar projections strongly vertical (fig. 7B, F), pygophore more elongate and with sinuous lateral margins (fig. 13D, K) 4
4. Eye shape in dorsal view distinctly greater than 90° angle (fig. 3A), corium uniformly brown (fig. 10E) E. obscura.
– Eye shape in dorsal view close to 90° angle (fig. 2A, E), corium with brown and yellow patterning (fig. 10A, B) 5
5. Yellow coloration on hemelytron largely restricted to proximal part of veins (fig. 10A), pygophore ventrally without transverse ridges along midline (fig. 13A) E. caballina.
– Yellow coloration on hemelytron on proximal part of veins and in adjacent cells (fig. 7B), pygophore ventrally with transverse ridges along midline (fig. 13H: pr) E. nouraguensis.
Ectrichodiella caballina, n. sp.
Figures 1A–C, 2A, B, 4A, B, 6A, B, 7A, B, 9A, C, 10A, 11A, B, 13A–G
Diagnosis. Recognized among species of Ectrichodiella by the large subapical spine on the scape (fig. 4B); eye in lateral view not reaching ventral head margin, about 0.5× height of head (fig. 2B); ocelli large and on paired elevations (fig. 2A); third labial segment 0.75× length of second (fig. 6A); large and strongly vertical midlateral scutellar projections (fig. 7B); proximal portion of M+Cu separated (fig. 10A), slightly inflated pterostigma (fig. 7A), yellow coloration on hemelytron largely restricted to proximal part of veins and narrow areas surrounding veins (fig. 10A); lateral margin of laterotergite 1+2 only slightly expanded (fig. 1A); posterior margin of tergum 7 indented (fig. 11A); posterior margin of sternum 8 slightly sinuous (fig. 13A), and pygophore ventrally without transverse ridges along midline.
Description. MALE: Macropterous, small body size. Measurements of holotype (in mm): habitus length from clypeus to abdominal margin: 3.74; head length excluding neck: 0.77; head greatest width: 0.98; synthlipsis width: 0.57; eye: 0.16; labial segment 2: 0.48; labial segment 3: 0.32; thorax length: 1.25; anterior pronotal lobe greatest length: 0.16; anterior pronotal lobe greatest width: 0.45; posterior pronotal lobe greatest length: 0.54; posterior pronotal lobe greatest width: 0.8; abdomen length: 1.97; abdomen greatest width: 1.84. COLORATION: Body brown; eye dark brown; ocellus pale; antennal setation light; thorax dark brown, with posterior pronotal lobe brown; legs yellow with brown band on femur; hemelytron with corium brown with yellow band proximally, with veins brown and yellow, yellow coloration on hemelytron largely restricted to proximal part of veins (fig. 10A) and distal margin of pterostigma; connexivum brown with contrasting yellow markings at sutures. VESTITURE: Head with sparse, thin, medium length yellow dorsally and with thin, short, white setae on ventral surface; labial setae thin, small, yellow; antennal setae less than 2× the diameter of antennal segment; thorax with sparse yellow, thin, medium length setae dorsally on anterior pronotal lobe and laterally on posterior lobe; scutellum with short yellowish setae densely lining lateral margins; small tubercles on hind trochanter; hemelytron with sparse, yellow, short, setae; connexivum with yellow short setae ventrally. STRUCTURE: HEAD: As in generic description, with interocular sulcus most pronounced at middle of head; synthlipsis 3× eye width (fig. 2A); with paired hemispherical elevations anterior of interocular sulcus; eye relatively large, height about 0.5× height of head (fig. 2B), reniform in lateral view, hemispherical in ventral view; ocelli large, on individual paired elevations. Antenna: as in generic description, with scape with large subapical spine (fig. 5B). Labium: as in generic description, with third segment 0.75× length of second (fig. 6A). THORAX: As in generic description, with surface smooth; anterior pronotal lobe less than half length and more than half width of posterior pronotal lobe, with anterior margin straight, with median longitudinal depression, with paired, rounded processes on disc (fig. 7B: alp) and paired glabrous areas on anterior pronotal lobe posterior to anterior lobe processes; posterior pronotal lobe with lateral ridges; midlateral scutellar projections large, strongly vertical (fig. 7B). Legs: as in generic description, with foretibia apically dilated (fig. 9A). Hemelytron: as in generic description, with distal margin of CuA1 cell fairly straight (fig. 10A), proximal portion of M+Cu separated (fig. 10A), and pterostigma slightly inflated. ABDOMEN: Lateral margin of laterotergite 1+2 slightly expanded; posterior margin of tergum 7 indented (fig. 11A); posterior margin of sternum 8 slightly sinuous (fig. 13A). Genitalia: pygophore with sinuous lateral margins (fig. 13A, B), ventral surface with distinct setation and ridges along midline (fig. 13A), posterior margin of pygophore slightly rounded (fig. 13H). Aedeagus as in figure 13E–G.
FEMALE: As in male, see table 1.
Etymology: The species is named after the Montagnes des Chevaux (“Horse Mountains”) in French Guiana where the holotype was collected, after the feminine form of the Latin adjective caballinus, referring to a horse.
Distribution: Only known from French Guiana.
Discussion: Ectrichodiella caballina, E. nouraguensis, and E. obscura, closely resemble each other with respect to overall habitus, color pattern, lateral ridges on the posterior pronotal lobe and size and vertical orientation of the midlateral scutellar projections. E. caballina, E. nouraguensis, and E. obscura, can be distinguished from each other by the wing color pattern, and details of the male genitalia among other features.
Holotype: Male: French Guiana: Guyane: Montagnes des Chevaux; 31 Jan 2010, SEAG, (UCR_ENT00127901) (MNMH).
Paratypes: French Guiana: Guyane: Montagnes des Chevaux: 28 Feb 2010, SEAG, 1 ♂(UCR_ENT00129410) (MNMH); 03 Jan 2010, SEAG, 1 ♂ (UCR_ENT00129411) (JMB); 21 Feb 2010, SEAG, 1 ♂ (UCR_ENT00129412) (UCR); 16 Jan 2009, SEAG, 1 ♂ (UCR_ ENT00129413) (MNMH); 09 Feb 2010, SEAG, 1 ♂ (UCR_ENT00129414) (UCR), 1 ♂ (UCR_ ENT00129417) (UCR); 28 Feb 2010, SEAG, 1 ♂ (UCR_ENT00129415) (UCR); 31 JAN 2010, SEAG, 1 ♂ (UCR_ENT00129416) (UCR); 24 Apr 2009; SEAG, 1 ♀ (UCR_ENT00129418) (UCR). Nouragues: 06 Mar 2011, SEAG, 1 ♀ (UCR_ENT00129404) (MNMH).
Ectrichodiella electrina, n. sp.
Figures 1G, H, 2C, D, 4C, D, 7C, D, 10C, D, 11C, D, 13O, P
Diagnosis: Recognized among species of Ectrichodiella by the large subapical spine on the scape (fig. 4D), ocelli large and on shared elevation (fig. 4C), without apical dilation of foretibia, proximal portion of M+Cu separated (fig. 10C, D), pterostigma strongly inflated (fig. 10C, D), posterior margin of tergum 7 slightly rounded (fig. 11C), sternum 8 posterior margin straight (fig. 13P), and pygophore posterior margin round (fig. 13O, P).
Description: MALE: Macropterous, small body size. Measurements of holotype (in mm): habitus length from clypeus to abdominal margin: 3.31; head length excluding neck: 0.41; head greatest width: 0.62; synthlipsis width: 0.34; eye: 0.12; labial segment 2 (1st visible): unknown; labial segment 3 (2nd visible): unknown; thorax length: 1.04; anterior pronotal lobe greatest length: 0.16; anterior pronotal lobe greatest width: 0.59; posterior pronotal lobe greatest length: 0.35; posterior pronotal lobe greatest width: 1.0; abdomen length: 1.77; abdomen greatest width: 1.26. COLORATION: Body brown; ocellus pale; antennal setation dark; thorax dark brown, with posterior pronotal lobe brown; legs dark; hemelytron with corium uniformly brown, with veins uniformly brown; connexivum uniformly dark brown. VESTITURE: Head fairly densely covered with short, curved setae; antennal setae less than 2× diameter of antennal segment; pronotum with short dark setae; scutellum with short setae densely lining lateral margins; small tubercles on hind trochanter not observable; connexivum with dark setae ventrally. STRUCTURE: HEAD: As in generic description, with interocular sulcus distinct throughout; with paired hemispherical elevations anterior of interocular sulcus; synthlipsis 2× eye width (fig. 2C); eye large, reaching ventral head margin; oval in lateral view, hemispherical in ventral view; ocelli large, on shared elevation (fig. 2C). Antenna: as in generic description, with scape with large subapical spine (fig. 4D). Labium: as in generic description, relative length of second and third segments not observable. THORAX: As in generic description, with surface slightly granulose; anterior pronotal lobe more than half length of and more than half width of posterior pronotal lobe, with anterior margin rounded, with median longitudinal depression, with paired, rounded processes on disc, but without glabrous areas posterior to processes; posterior pronotal lobe with lateral ridges (fig. 7C); midlateral scutellar projections large, mostly horizontal. Legs: as in generic description, without dilation apically on foretibia apically; ventroapical projections of forefemur and foretibial comb not observable (fig. 9D). Hemelytron: as in generic description, with distal margin of CuA1 cell fairly straight (fig. 10D), proximal portion of M+Cu separated, and pterostigma strongly inflated (fig. 10C). ABDOMEN: Lateral margin of laterotergite 1+2 slightly expanded; posterior margin of tergum 7 slightly rounded (fig. 11D); posterior margin of sternum 8 straight. Genitalia: pygophore with rounded lateral margins (fig. 13O, P), ventral surface without distinct setation and ridges along midline (fig. 13P), posterior margin of pygophore distinctly rounded (fig. 13H). Aedeagus not observed.
FEMALE: Unknown.
Etymology: The species name is the feminine form of the Neo-Latin adjective electricus meaning “of amber” and referring to the type of fossilization.
Distribution: Dominican Republic, Dominican amber.
Discussion: Ectrichodiella electrina is unambiguously classified as part of Ectrichodiella shown by the morphology-based phylogenetic analysis in this study. Ectrichodiella electrina was recovered as sister taxon to either E. minima and E. rafaeli or the clade formed by E. caballina, E. nouraguensis, and E. obscura. Ectrichodiella electrina is unique among species of Ectrichodiella in the absence of the apical dilation of the foretibia and the pygophore posterior margin being round. While this specimen is excellently preserved, certain features are difficult to observe (e.g., the head in lateral view) or document (e.g., details of the labium; wing slightly twisted). Overall, the documentation of morphological features is sufficient to place this species in the genus Ectrichodiella and to distinguish it from all other known congeners.
Holotype: Male: Dominican Republic; Dominican Amber specimen, Oligo-Miocene (AMNH_DR-14-405) (AMNH).
Ectrichodiella minima (Valdés, 1910)
Ectrichodia minima Uhler (manuscript name).
Ectoiocchoda [sic] minima Valdés, 1910.
Ectrichodiella cubensis Fracker and Bruner, 1924; junior synonym: Bruner and Barber (1937).
Ectrichodiella minima Bruner and Barber, 1937.
Revised diagnosis: Recognized among species of Ectrichodiella by the small subapical spine on scape, ocelli large and on individual paired elevations, the posterior pronotal lobe without lateral ridges, midlateral scutellar projections small, mostly horizontal, the proximal portion of M+Cu fused, lateral margin of laterotergite 1+2 with small spine, and the pygophore posterior margin distinctly round.
Redescription: MALE: Macropterous, small body size. Dimensions (in mm; from Gil-Santana et al., 2013): habitus length from clypeus to abdominal margin: 3.39; head length: 0.41; head width: 0.62; synthlipsis width: 0.38; labial segment 2 (1st visible): 0.51; labial segment 3 (2nd visible): 0.38; anterior pronotal lobe length: 0.23; anterior pronotal lobe width: 0.63; posterior pronotal lobe length: 0.51; posterior pronotal lobe width: 1.19; abdomen greatest width: 1.79. COLORATION: Body yellow; eye brown; antennal setae light; thorax light brown; legs brown; hemelytron with corium uniformly brown, with veins uniformly yellow; connexivum uniformly dark yellow. VESTITURE: Head covered with fringe of short white setae; antenna setae greater than 2× the diameter of antenna segment; thorax with fringe short whitish setae on anterior margin, sutures covered by dense fringe of short white pubescence; pronotum with short, dense, pubescence; scutellum with long thin yellowish setae; small tubercles on hind trochanter; hemelytron with veins and cuneal area with sparse long, thin yellow setae; abdomen with long setae. STRUCTURE: HEAD: As in generic description, with interocular sulcus most pronounced anterior to ocelli; with paired hemispherical elevations anterior of interocular sulcus; synthlipsis more than 3× eye width; eye large, reniform in lateral view; ocelli large, on individual paired elevations. Antenna: as in generic description, with scape with small subapical spine. Labium: as in generic description, with third segment 0.75× length of second. THORAX: As in generic description, with surface smooth; anterior lobe more than half length and less than half width of posterior lobe, with anterior margin rounded, with median longitudinal depression, with paired, rounded processes on disc and without pair of glabrous areas posterior to processes; posterior pronotal lobe without lateral ridges; midlateral scutellar projections small, mostly horizontal. Legs: as in generic description, with foretibia apically dilated. Hemelytron: as in generic description, with distal margin of CuA1 cell fairly straight, proximal portion of M+Cu fused, and pterostigma strongly inflated. ABDOMEN: Lateral margin of laterotergite 1+2 with small spine; posterior margin of tergum 7 straight; sternum 8 not observed. Genitalia (based on Gil-Santana et al., 2013: plate 3): pygophore with rounded lateral margins, ventral surface without distinct setation and ridges along midline, posterior margin distinctly rounded. Aedeagus simple and slender.
FEMALE: As in male.
Distribution: Cuba.
Discussion: The taxonomic history of this species was outlined in Gil-Santana et al. (2013). That publication also provides more details on the morphology of this species, in particular vestiture, coloration, additional measurements, as well as habitus photographs and genitalic illustrations. Our morphology-based phylogenetic analysis recovered this species from the Greater Antilles as sister species to E. rafaeli that is known from French Guiana and the Amazon basin.
Material examined: CUBA: Santiago: Palma Peak, Sierra Maestra, 1386 m, 10 Jul 1922 - 20 Jul 1922, S.C. Bruner and C.H. Ballou, Holotype of Ectrichodiella cubensis, 1 ♀ (UCR_ENT 00008147) (USNM).
FIG. 16.
Phylogenetic hypothesis of Ectrichodiella based on the six ingroup species, five outgroup taxa, and 45 morphological characters. The tree shown is the best tree derived from the implied weights analysis after jackknifing. Values shown underneath branches are jackknife values <50% with values from IW on left and EW on right. The fossil species E. electrina, is here shown as sister to the two described species of Ectrichodiella, although the node lacks support.
Ectrichodiella nouraguensis, n. sp.
Figures 1D–F, 2E, F, 4E, F, 6C, D, 7E, F, 9B, D, 10B, 11E, F, 13H–N
Diagnosis: Recognized among species of Ectrichodiella by the large subapical spine on scape (fig. 4E), eye in lateral view not reaching ventral head margin, about 0.6× height of head (fig. 2F), ocelli large, on individual elevations (fig. 2E), third labial segment 0.5× length of second (fig. 6C), large and strongly vertical midlateral scutellar projections (fig. 7F), proximal portion of M+Cu separated (fig. 10B), slightly inflated pterostigma (fig. 10B), yellow coloration on hemelytron on proximal part of veins and in adjacent cells (fig. 10B), lateral margin of laterotergite 1+2 only slightly expanded (fig. 1D), posterior margin of tergum 7 straight (fig. 11E), and posterior margin of sternum 8 strongly sinuous (fig. 13H), and pygophore ventral surface with distinct setation and ridges along midline (fig. 13H).
Description: MALE: Macropterous, small body size. Dimensions (in mm): habitus length from clypeus to abdominal margin: 3.32; head length excluding neck: 0.52; head greatest width: 0.63; synthlipsis width: 0.49; eye: 0.16; labial segment 2 (1st visible): 0.48; labial segment 3 (2nd visible): 0.35; thorax length: 1.12; anterior pronotal lobe greatest length: 0.16; anterior pronotal lobe greatest width: 0.8; posterior pronotal lobe greatest length: 0.44; posterior pronotal lobe greatest width: 1.5; abdomen length: 1.68; greatest width at abdomen: 1.6. COLORATION: Body dark brown; eye dark brown; antennal setation light; thorax dark brown with posterior pronotal lobe brown; legs yellow with band of brown on femur, hemelytron with corium brown with yellow band proximally with veins brown and yellow, yellow coloration on hemelytron on proximal part of veins and in adjacent cells (fig. 10B); connexivum brown with contrasting yellow markings at sutures. VESTITURE: Head with light yellow-colored setae, with fringe of white setae behind eyes, labium setae small, yellow, thin; antenna setae greater than 2× the diameter of the antenna segment; pronotum sparse short yellow setae laterally; thorax with short, fine, yellowish setae ventrally; scutellum long thin yellow setae along sides; small tubercles on hind trochanter; hemelytron with sparse, short, yellow, setae; connexivum with yellow short setae on ventral side. STRUCTURE: HEAD: As in generic description, with interocular sulcus most pronounced at middle of head; with paired hemispherical elevations anterior of interocular sulcus; synthlipsis 2× width of eye (fig. 2E); eye relatively large, about 0.6× height of head, reniform in lateral view; ocelli large, on individual paired elevations (fig. 2E). Antenna: as in generic description, with scape with large subapical spine (fig. 4F). Labium: as in generic description, with third segment 0.5× length of second segment (fig. 6C). THORAX: As in generic description, with surface smooth; anterior pronotal lobe less than half length and more than half width of posterior pronotal lobe, with anterior margin rounded, with median longitudinal depression, with paired, rounded process on disc and with pair of glabrous areas posterior to processes; posterior pronotal lobe with lateral ridges (fig. 7E); midlateral scutellar projections large, strongly vertical (fig. 7F). Legs: as in generic description, with foretibia apically dilated. (fig. 9A: td). Hemelytron: as in generic description, with distal margin of CuA1 cell fairly straight, proximal portion of M+Cu separated, and pterostigma slightly inflated (fig. 10B). ABDOMEN: Lateral margin of laterotergite 1+2 slightly expanded; posterior margin of tergum 7 straight (fig. 11E); posterior margin of sternum 8 strongly sinuous. Genitalia: pygophore with sinuous lateral margins (fig. 13H–J), ventral surface with distinct setation and ridges along midline (fig. 13H), posterior margin of pygophore slightly rounded (fig. 13I). Aedeagus as in figure 13L–N.
FEMALE: As in male, see table 1.
Etymology: The species is named for Nouragues Nature Reserve in French Guiana where the holotype was collected.
Distribution: Currently only known from French Guiana.
Discussion: See discussion under E. caballina.
Holotype: Male: French Guiana: Cayenne: Regina: Pararé, Nouragues Nat. Res., 4.03800, -52.67286; 70 m (230 ft), 01 Nov 2009, SEAG, (UCR_ENT00127900) (MNMH).
Paratypes: French Guiana: Guyane: Nouragues: 05 Feb 2010, SEAG, 1 ♂ (UCR_ ENT00129397) (UCR); 01 Nov 2009, SEAG, 1 ♂ (UCR_ENT00129398) (MNMH); 28 Jan 2010, SEAG, 1 ♂ (UCR_ENT00129400) (UCR); 05 Feb 2010, SEAG, 1 ♂ (UCR_ENT00129401) (MNMH); 30 Nov 2009, SEAG, 1 ♂ (UCR_ENT00129402) (UCR); 05 Feb 2010, SEAG, 1 ♂(UCR_ENT00129403) (UCR); 30 Nov 2009, SEAG, 1 ♂ (UCR_ENT00129419) (UCR). Itoupe: 17-30 Mar 2010, SEAG, 1 ♂ (UCR_ENT00129422) (JMB). Saul: 13 Aug 2010, SEAG, 1 ♀(UCR_ENT00129405) (UCR). Montagnes des Chevaux: 31 JAN 2010, SEAG, 1 ♀ (UCR_ ENT00129406) (MNMH).
Ectrichodiella obscura, n. sp.
Figures 1I–K, 3A, B, 5A, B, 6, E, F, 8A, B, 10E, 12A, B, 14A–G
Diagnosis: Recognized among species of Ectrichodiella by the large subapical spine on scape (fig. 5A), eye in lateral view not reaching ventral head margin, about 1× height of head (fig. 3A), ocelli large, on individual elevations (fig. 3B), third labial segment 0.5× length of second (fig. 6F), large strongly vertical midlateral scutellar projections (fig. 8B), proximal position of M+Cu separated, slightly inflated pterostigma (fig. 10E), hemelytron corium dark brown with veins brown and yellow, lateral margin on laterotergite 1+2 only slightly expanded (fig. 11J), posterior margin of tergum 7 straight (fig. 12A), and posterior margin of sternum 8 slightly sinuous (fig. 13Q), and pygophore ventral surface with distinct setation along midline.
Description: MALE: Macropterous, small body size. Dimensions (in mm): habitus length from clypeus to abdominal margin: 4.83; head length excluding neck: 0.67; head greatest width: 0.94; synthlipsis width: 0.5; eye: 0.19; labial segment 2 (1st visible): 0.69; labial segment 3 (2nd visible): 0.39; thorax length: 0.98; anterior pronotal lobe greatest length: 0.24; anterior pronotal lobe greatest width: 0.8; posterior pronotal lobe greatest length: 0.72; posterior pronotal lobe greatest width: 1.42; abdomen length: 2.7; greatest width at abdomen: 2.2. COLORATION: Body dark brown (fig. 1I); eye dark brown; ocellus pale; antennal setation light; thorax dark brown, with posterior pronotal lobe dark brown; legs yellow with brown band on femur; hemelytron with corium dark brown with veins brown and yellow; connexivum dark brown with contrasting yellow markings at sutures (fig. 1J). VESTITURE: Head with light yellow colored hair, with fringe of hair covering posterior surface of head (fig. 3B); labial setae thin, small, yellow; antennal setae greater than 2× the diameter of antennal segment (fig. 5A); thorax with sparse yellow, thin, short setae dorsally on anterior pronotal lobe and laterally on posterior lobe; scutellum with dense short yellow hairs along sides; small tubercles on hind trochanter; hemelytron with yellow, short, setae along the Sc vein; connexivum with yellow short setae ventrally. STRUCTURE: HEAD: As in generic description, with interocular sulcus most pronounced at ocelli; synthlipsis 2× eye width (fig. 3A); with paired hemispherical elevations anterior of interocular sulcus; eye relatively large, height about 0.5 height of head, reniform in lateral view, quarter spherical in ventral view; ocelli large, on individual paired elevations (fig. 3B). Antenna: as in generic description, with scape with large subapical spine (fig. 5B). Labium: as in generic description, with third segment 0.5× length of second (fig. 6A). THORAX: As in generic description, with surface smooth; anterior pronotal lobe less than half length and more than half width of posterior pronotal lobe, with anterior margin rounded, with median longitudinal depression, with paired, rounded processes on disc and with pair of glabrous areas on anterior pronotal lobe posterior to anterior lobe processes; posterior pronotal lobe with lateral ridges (fig. 8A); midlateral scutellar projections large, strongly vertical (fig. 8B). Legs: as in generic description, with foretibia apically dilated. Hemelytron: as in generic description, with distal margin of CuA1 cell fairly straight, proximal portion of M+Cu separated, and pterostigma slightly inflated (fig. 10E). ABDOMEN: Lateral margin of laterotergite 1+2 slightly expanded (fig. 12A); posterior margin of tergum 7 straight; posterior margin of sternum 8 slightly sinuous (fig. 14A). Genitalia: pygophore with sinuous lateral margins, ventral surface with distinct setation along midline, posterior margin of pygophore rounded (fig. 14B–D). Aedeagus as in figure 14E–G.
Female: As in male, see table 1.
Etymology: The species is named for its dark coloration.
Distribution: Currently only known from French Guiana.
Discussion: See discussion under E. caballina.
Holotype: Male: French Guiana: Guyane: Saul: 23 Feb 2011, SEAG, (UCR_ENT00129408) (MNMH).
Paratypes: French Guiana: Guyane: Saul belvedere: 3.73348, -53.36236; 17 Jan 2011, SEAG, 1 ♂ (UCR_ENT00129431) (UCR); 24 Jan 2011, SEAG, 1 ♂ (UCR_ENT00129432) (JMB); 11 Jan 2011, SEAG, 1 ♂ (UCR_ENT00129433) (UCR). Saul: 13 May 2011, SEAG, 1 ♂(UCR_ENT00129434) (UCR); 31 Jan 2011, SEAG, 1 ♂ (UCR_ENT00129435) (UCR). Montagnes des Chevaux: 01 Nov 2011, SEAG, 1 ♂ (UCR_ENT00129709) (UCR). Nouragues: 23 Feb 2011, SEAG, 1 ♀ (UCR_ENT00129429) (MNMH).
Ectrichodiella rafaeli (Gil-Santana and Coletto-Silva, 2005)
Figures 1L–N, 3C, D, 5C, D, 6G, H, 8C, D, 10F, 12C, D, 14H–N
Berengeria rafaeli Gil-Santana and Coletto Silva, 2005: 35.
Revised diagnosis: Recognized among species of Ectrichodiella by the absence of a subapical spine on the scape, small eyes, the ocelli on flat dorsal head surface, the posterior pronotal lobe with lateral shallow ridges only on basal half, the proximal portion of M+Cu fused, and the lateral margin of laterotergite 1+2 with a large, curved spine, posterior margin of sternum 8 slightly sinuous, and pygophore ventral surface with distinct setation.
Redescription: FEMALE: Macropterous, small body size. Dimensions (in mm; from Gil-Santana et al., 2013): habitus length from clypeus to abdominal margin: 3.55; head length: 0.65; head width: 0.63; synthlipsis width: 0.45; labial segment 2 (1st visible): 0.45; labial segment 3 (2nd visible): 0.2; anterior pronotal lobe length: 0.3; anterior pronotal lobe width: 0.65; posterior pronotal lobe length: 0.45; posterior pronotal lobe width: 1.2; abdomen length: 1.7; abdomen greatest width: 1.7. COLORATION: Body brownish (fig. 1L–N); eye dark; antennal setae light; thorax red brown, with posterior pronotal lobe red brown; legs reddish; hemelytron with corium generally brown, with veins basely and their surrounding areas pale, almost yellowish; connexivum uniformly dark yellow, with sutures white. VESTITURE: Head covered posteriorly and ventrally with white fringe setae (fig. 3C); labium with short, light colored setae; antenna setae greater than 2× the diameter of the antenna segment; thorax covered with long, fine, yellowish setae and short, whitish setae on anterior margin; pronotum brown with many long, thin setae; scutellum with long and dense, short hairs along edges of scutellum (fig. 8C); small tubercles on hind trochanter; hemelytron with sparse fine setae; abdomen with first visible sternite (sternite 2) with fringe of short white setae, others with sparse long bright setae; connexivum with long bright setae mainly on margin. STRUCTURE: HEAD: As in generic description, without paired hemispherical elevations anterior of interocular sulcus (fig. 3C); synthlipsis more than 3× eye width; eye small, reniform in lateral view, hemispherical in ventral view; ocelli small, on flat dorsal head surface (fig. 3D). Antenna: as in generic description, with small subapical spine on scape (fig. 5D). Labium: as in generic description, with third segment 0.5× length of second segment (fig. 6G). THORAX: As in generic description, with surface granulose on lateral sides of anterior lobe; anterior pronotal lobe more than half length and less than half width of posterior lobe, with anterior margin rounded, median longitudinal depression created by a series of small shallow subcircular impressions (fig. 8C), greatest at midpoint and absent posteriorly, with a small spine on left side of lateral margin at posterior half, and anterior pronotal lobe processes conspicuous, spiniform (documented in Gil-Santana et al., 2013 fig. 28); posterior pronotal lobe with lateral shallow ridges on basal half; midlateral scutellar projections small, mostly horizontal (fig. 8D). Legs: as in generic description, with foretibia apically dilated. Hemelytron: as in generic description, with distal margin of CuA1 cell slightly curved, proximal portion of M+Cu fused, and pterostigma strongly inflated (fig. 10F). ABDOMEN: Lateral margin of laterotergite 1+2 with large, curved spine (fig. 12C). Genitalia: documented by Gil-Santana et al. (2013: fig. 35).
MALE: As in female, see Table 1. ABDOMEN: posterior margin of tergum 7 straight (fig. 12C); posterior margin of sternum 8 slightly sinuous. Genitalia: pygophore with rounded lateral margins (fig. I–K), ventral surface with distinct setation, posterior margin of pygophore rounded. Aedeagus as in figure 14 L–N.
Distribution: Brazil and French Guiana.
Discussion: This species was described in a new genus, Berengeria, in the subfamily Reduviinae by Gil-Santana and Coletto-Silva (2005), in part owing to features found in other genera of this subfamily such as described by Gil-Santana et al. (2013):
two-segmented tarsi, absence of fossula spongiosa in tibiae, and two cells in the membrane of hemelytra (Nalata Stål, 1860); a relative reduction of the area of the corium when compared to the membrane of hemelytra (Microlestria Stål, 1872) as well as the general resemblance, hemelytral coloration, and long body hairs like Peregrinator biannulipes (Montrouzier and Signoret, 1861) .
Berengeria was synonymized with Ectrichodiella by Gil-Santana et al. (2013). Those authors also provide additional morphological descriptions, in particular detailed accounts of the vestiture, coloration, and additional measurements. Our morphology-based phylogenetic analysis recovered this species from Brazil as sister species to E. minima from the Greater Antilles.
Material examined: Only photographs of the female holotype were seen. Collection event information in Gil-Santana et al. (2013): BRAZIL, Amazonas, Reserva Ducke, 26 km N of Manaus, 27-VIII-1982/ J. A. Rafael [leg.], Ar. Malaise / Hemipt 155 Holotipo / HOLOTIPO, Berengeria rafaeli Gil-Santana and Coletto-Silva, 2005 (red label) [INPA]. French Guiana: Guyane: Nouragues: 30 Nov 2009; SEAG, 1 ♂ (UCR_ENT00129344) (UCR); 08 Jun 2010, SEAG, 1 ♂ (UCR_ENT00129421) (MNMH). Montagnes des Chevaux: 16 Oct 2010, SEAG, 1 ♂ (UCR_ENT00129407) (MNMH); 16 Dec 2009, SEAG, 1 ♀ (UCR_ENT00129409) (UCR). Saul: 31 Jan 2011, SEAG, 1 ♂ (UCR_ENT00129420) (UCR).
ACKNOWLEDGMENTS
We are grateful to David A. Grimaldi for making available the holotype of the uniquely well-preserved Dominican amber fossil that is part of this study. We also thank the Société Entomologique Antilles-Guyane for their efforts in uncovering the insect biodiversity of French Guiana. We acknowledge the Willi Hennig Society for making TNT widely available. For financial support, we thank the University of California, Riverside (research support and graduate student assistantship support to T.B.). For comments on earlier drafts of this study, we thank former and current members of the Weirauch lab.
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