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Though the phylogenetic signal of loci on sex chromosomes can differ from those on autosomes, chromosomal-level genome assemblies for non-vertebrates are still relatively scarce and conservation of chromosomal gene content across deep phylogenetic scales has therefore remained largely unexplored. We here assemble a uniquely large and diverse set of samples (17 Anchored Hybrid Enrichment [AHE], 24 RNA-Seq, and 70 whole-genome sequencing [WGS] samples of variable depth) for the medically important assassin bugs (Reduvioidea). We assess the performance of genes based on multiple features (e.g., nucleotide vs. amino acid, nuclear vs. mitochondrial, and autosomal vs. X chromosomal) and employ different methods (concatenation and coalescence analyses) to reconstruct the unresolved phylogeny of this diverse (∼7,000 spp.) and old (>180 MYA) group. Our results show that genes on the X chromosome are more likely to have discordant phylogenies than those on autosomes. We find that the X chromosome conflict is driven by high gene substitution rates that impact accuracy of phylogenetic inference. However, gene tree clustering showed strong conflict even after discounting variable third codon positions. Alternative topologies were not particularly enriched for sex chromosome loci, but spread across the genome. We conclude that binning genes to autosomal or sex chromosomes may result in a more accurate picture of the complex evolutionary history of a clade.

Dipsocoromorpha, or the minute litter bugs, are a poorly studied and minuscule group of true bugs (Hemiptera: Heteroptera) with uncertain phylogenetic position, often bizarre morphology, and substantial undescribed biodiversity. A combination of taxonomic revisions, comparative morphological studies, and phylogenetic analyses based on both morphological and molecular data is employed to advance our knowledge of the group. The first chapter taxonomically revises the New World genus Chinannus and describes 26 new species. The second chapter develops and tests a cost-efficient DNA sequencing method for archival specimens. The third chapter uses and refines this method to conduct a comprehensive phylogenetic analysis of a very diverse group of dipsocoromorphans, the Corixidea genus group. The fourth chapter taxonomically revises the genus Voragocoris that belongs to the Corixidea genus group, building on the phylogeny inferred in chapter three, and describing seven new species. The fifth chapter conducts a comparative study of abdominal morphology in Dipsocoromorpha, and both standardizes the terminology and proposes primary homology hypotheses, that could be used for a phylogenetic reconstruction. The sixth, and last, chapter presents a comprehensive phylogenetic analysis of Dipsocoromorpha based on both morphological and molecular data.

Currently comprising only about 430 species, Dipsocoromorpha or minute litter bugs are one of the small infraorders of Heteroptera. They are classified into five morphologically distinct families--Ceratocombidae, Dipsocoridae, Hypsipterygidae, Schizopteridae and Stemmocryptidae--but relationships among and within these families are poorly understood owing to the lack of phylogenetic studies. A phylogenetic hypothesis based on combined molecular and morphological data is important to both evaluate and revise the higher-level classification and to explore the evolutionary history of morphological features including elytriform (or beetle-like) forewings, that seem to have evolved multiple times in this group, jumping devices, and the highly modified and diverse male genitalia. We here use a dataset combining Sanger-derived (~4500 bp; 108 taxa) and Illumina-generated (~7500 bp; 24 taxa) sequence data with a morphological matrix (159 characters) and taxon sampling that comprises all currently recognized family-, subfamily- and tribal-level taxa and comprehensive genus-level sampling to investigate phylogenetic relationships within litter bugs. Our results support the monophyly of Dipsocoromorpha, Schizopteridae and Dipsocoridae, whereas Ceratocombidae and the schizopterine subfamily "Ogeriinae" are polyphyletic and paraphyletic, respectively. A new classification is proposed that recognizes six families, including Trichotonannidae, stat.n., and two subfamilies each within the two larger families Ceratocombidae and Schizopteridae. Ancestral state reconstructions outline the complex evolutionary history of many morphological characters, including 15 independent origins of elytriform forewings, and at least five shifts in the degree of genitalic asymmetry.

The position of some taxa on the Tree of Life remains controversial despite the increase in genomic data used to infer phylogenies. While analyzing large datasets alleviates stochastic errors, it does not prevent systematic errors in inference, caused by both biological (e.g., incomplete lineage sorting, hybridization) and methodological (e.g., incorrect modeling, erroneous orthology assessments) factors. In our study, we systematically investigated factors that could result in these controversies, using the treeshrew (Scandentia, Mammalia) as a study case. Recent studies have narrowed the phylogenetic position of treeshrews to three competing hypotheses: sister to primates and flying lemurs (Primatomorpha), sister to rodents and lagomorphs (Glires), or sister to a clade comprising all of these. We sampled 50 mammal species including three treeshrews, a selection of taxa from the potential sister groups, and outgroups. Using a large diverse set of loci, we assessed support for the alternative phylogenetic position of treeshrews. A plurality of loci support treeshrews as sister to rodents and lagomorphs; however, only a few loci exhibit strong support for any hypothesis. Surprisingly, we found that a subset of loci that strongly support the monophyly of Primates, support treeshrews as sister to primates and flying lemurs. The overall small magnitude of differences in phylogenetic signal among the alternative hypotheses suggests that these three groups diversified nearly simultaneously. However, with our large dataset and approach to examining support, we provide evidence for the hypothesis of treeshrews as sister to rodents and lagomorphs, while demonstrating why support for alternate hypotheses has been seen in prior work. We also suggest that locus selection can unwittingly bias results.

Guapinannus uncus, sp. n. (Figs 4, 7, 9) Holotype. Male. BELIZE: Belize, 17.18988°N 88.49765°W, 55m, no date provided, collector unknown, 1♂ (UCR _ ENT 00090858) (FMNH). Paratypes: BELIZE: Orange Walk: Rio Bravo Cons. Area 3 Well Tran (Near Res. Station), 18.06667°N 88.55°W, 10m, 10 Sep 1995, P. W. Kovarik, 1♂ (UCR _ ENT 00094283) (UCR). Etymology: Named for the hook-shaped vesica after “uncus” Latin for hook (noun in apposition). Diagnosis: Recognized by the small mid-sized body length (0.67 mm), elongate oval body shape (length to width ratio 3.4) (Fig. 4), and apex of vesica hook-shaped (Fig. 7). Description: Male: Total body length 1.19 mm; length from posterior margin of pronotum to posterior wing margin 0.97 mm; greatest width across pronotum 0.27 mm; greatest width across forewings 0.35 mm. General coloration: uniformly pale. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars indistinct. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum small, evenly distributed; scutellum in dorsal view gradually narrowed to tip, tip pointed. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin rounded, vein-tracing areoles on proximal part of wing absent; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium inflated, elongate, with median notch, without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc slightly wider than other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2–3 trapezoidal; cub almost reaching wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, surpassing right pygophore margin, thick at midpoint, with bottle opener-shaped apex, close to apex rounded; anophoric ridge with process, process short, with single lobe curved, tip pointed; right paramere curved, broad throughout, basal process broad; left paramere straight, tapering, basal process broad. Female: Unknown. Collecting method and habitat: Collected using flight intercept trap.

Guapinannus anaticulus, sp. n. (Figs 2, 7, 9) Holotype. Male. GUATEMALA: San Marcos: Bojonal Road, 14.94786°N 91.8837°W, 1618 m, 04 Jun 2015, R. Anderson, 1♂ (UCR _ ENT 00127123) (UCR). Etymology: Named for the duck head-shaped basal process of the right paramere after “anas” meaning duck in Latin. Diagnosis: Recognized by the mid-sized body length (1.45 mm), oval forewing outline, narrow C+SC (black arrow), rc1 about 3 times as long as wide and slightly wider anteriorly (white arrow), tc more than 3 times as long as wide (Fig. 2), the short, slender, and straight vesica with blunt, expanded tip, and the broad, duck head-shaped basal process of the right paramere (Fig. 7). Description: Male: Total body length 1.45 mm; length from posterior margin of pronotum to posterior wing margin 1.23 mm; greatest width across pronotum 0.63 mm; greatest width across forewings 0.81 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long; setae on posterior margin of forewing extremely long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum mediumsized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip pointed. Forewing: With outline oval; membrane contributing about one third of forewing length, posterior distal margin rounded, vein-tracing areoles on proximal part of wing absent; Sc+C uniform, relatively narrow; distal process of R pronounced; An1 wing organ on corium strongly inflated, rounded, with median notch, without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 5 times as long as wide; tc more than 3 times as long as wide; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 almost triangular; cub almost reaching wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, surpassing right pygophore margin, thick at midpoint, tapering to blunt, expanded tip, close to apex slightly s-shaped; anophoric ridge with process, process short, with single lobe curved, tip blunt; right paramere curved, tapering, basal process duck head-shaped; left paramere straight, tapering, basal process broad. Female: Unknown. Collecting method and habitat: Collected in cloud forest litter using sifting leaf litter., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on pages 266-267, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718

Guapinannus Wygodzinsky, 1951 (Figs 1–9) Type species: Guapinannus bierigi Wygodzinsky, 1951 Diagnosis: Recognized among Schizopteridae by the relatively small eyes (Fig. 5C, D), four-segmented labium (Fig. 1A), flat and wide forewings (Figs 1–4), unique forewing venation with long fracture in the costal margin, long rc1, rc, and tc cells (Figs 1B, 2, 6A), C+S, with An1/2 claval process except G. dispar sp. n. (Figs 2, 6A, D), better developed in males than in females (Fig 1B), males with wing organ associated with An1 on corium adjacent to the claval process (Figs 2–4, 6A, D), 2-2-3 tarsal formula in both sexes, male genitalia simple (Fig. 7) and female with well-developed ovipositor (Figs 1C, 8). Re-description: Male: Ovoid (greatest length to width ratio 1.57–1.85) to elongate ovoid (length to width ratio 3.4), total body length 0.66–2.06 mm; length from posterior margin of pronotum to posterior wing margin (0.51–1.49 mm). General coloration: uniformly pale or a mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long or short, setae on posterior margin of forewing long or extremely long. Structure: Head: Eye ranging from small (less than one quarter head height) to relatively large (almost half as high as head) (Fig. 5C, D); ocellus adjacent to eye (Fig. 5D), at 11 o’clock position seen in lateral view and about twice the size of one ommatidium; four-segmented labium with segments 1, 2 and 3 of similar length and 4 about 2.5–3 times as long as any other segment (Fig. 1A); muscle scars present on the frons and vertex present (Figs 1A, 5C). Thorax: Collar in dorsal view narrow (Fig. 5C); proepisternal lobe inflated (Fig. 5D); separation of anterior and posterior lobes (dorsal view) marked by weak depression (Fig. 5C); anterior pronotal lobe with muscle scars, margin concave or straight; posterior margin slightly concave to almost straight; pits on pronotum ranging from small to large, evenly distributed or grouped (Fig. 1A); scutellum in dorsal view abruptly or gradually narrowed to tip, tip pointed, rounded, or slightly inflated (Figs 2–4). Forewing: With outline elongate oval or broad anteriorly; with (G. dispar, sp. n., Fig. 2) or without (all other species) distinct honey-comb pattern; membrane (cub, dc1, rc2–3 and areas distal to these cells) contributing about one third or less to forewing length; posterior distal margin rounded or square; clavus twice as long as wide; vein-tracing areoles along margins of a combination of scc, rc1, rc1, rc2, and tc on proximal part of wing absent or present (Fig. 6A, C); C+Sc narrow (Fig. 2, G. anaticulus, sp. n.), fairly broad (Fig. 2, G. clava, sp. n.), broad (Fig. 2, G. falcis, sp. n.), or very broad (Fig. 3, G. lutuosus, sp. n.), either of uniform width (Fig. 2, G. anaticulus, sp. n.) or wider at widest part of wing (Fig. 2, G. auriculu s, sp. n.); R flattened in distal part (black arrow in Fig. 1B); distal process of R pronounced (white arrow in Fig. 1B) or weakly developed; An1/2 claval process well developed in males (except G. tenuis, sp. n.), overlapping base of An1 wing organ, apex acute and in lateral view pointing dorsad (Figs 2–4, 6A, D); An1 wing organ on corium strongly inflated in all males, with or without (e.g., Fig. 4, G. tergus, sp. n.) notch articulating with claval process, wing organ rounded, elongate, or acute, with or without median notch; scc about 1.5 times longer than wide (Fig. 6A) (except ~2 times longer than wide in G. castigatus, sp. n.); scc undivided except in G. dispar, sp. n. (Fig. 2); rc1 about 3 times as long as wide except in G. dispar, sp. n., ranging from parallel-sided to slightly or much wider anteriorly; rc about 4 or 5 times as long as wide; tc 3 times or more than 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins (e.g., Fig. 2, G. artus, sp. n.) or thicker (e.g., Fig. 2, G. falcis, sp. n.); distal margin of rc1, rc, and tc cells double s-shaped (Fig. 2, G. anaticulus, sp. n.), s-shaped (Fig. 1, G. bierigi), or angular s-shaped (Fig. 2, G. dispar, sp. n.); rc2–3 almost triangular (e.g., Fig. 2, G. anaticulus, sp. n.), trapezoidal (e.g., Fig. 2, G. castigatus, sp. n.), or rectangular (Fig. 2, G. dispar, sp. n.); cub oval, almost reaching wing margin or distant form wing margin (except elongate in G. dispar; Fig. 2); dc1 with basal portion fairly narrow and elongate (Fig. 2, G. anaticulus, sp. n.) or relatively wide and short (Fig. 2, G. auriculu s, sp. n.); M beyond Cu short. Legs: males and females with tarsal formula 2-2-3, with claws long except short in G dispar, sp. n. Genitalia (Fig. 7): Pygophore without lobe on right side (Fig. 7, G. anaticulus, sp. n.), with small (Fig. 7, G. artus, sp. n.) or large lobe (Fig. 7, G. auriculus, sp. n.), or with slightly extended right margin (Fig. 7, G. castigatus, sp. n.). Vesica about one half loop (e.g., Fig. 7, G. castigatus, sp. n.), between half and one loop (e.g., Fig. 7, G. anaticulus, sp. n.), or one loop (e.g., Fig. 7, G. plurilobus, sp. n.), either not reaching right pygophore margin or surpassing margin, at midpoint thick (e.g., Fig. 7, G. graziae, sp. n.), very thick (e.g., Fig. 7, G. robustus, sp. n.), or more slender (e.g., Fig. 7, G. falcis, sp. n.), tapering to blunt or acute tip, tip itself blunt, bottle-opener shaped (e.g., Fig. 7, G. uncus, sp. n.), slightly s-shaped (e.g., Fig. 7, G. falcis, sp. n.), rounded (e.g., Fig. 7, G. artus, sp. n.), or fairly straight (e.g., Fig. 7, G. tatumbia, sp. n.). Anophoric ridge with process present (e.g., Fig. 7, G. artus, sp. n.) or absent (Fig. 7, G. castigatus, sp. n.), process short or long, with single lobe curved or relatively straight, or with multiple lobes, tip pointed, blunt, or combination of pointed and blunt. Right paramere curved or straight, tapering towards apex or broad throughout; basal process of paramere broad, narrow, thumb-shaped, or duck head-shaped. Left paramere straight or curved, tapering towards apex or broad throughout, basal process broad or narrow. Female: As male, but with less well developed An1/2 claval process and without An1 wing organ on corium (Fig. 1B). Genitalia: with well-developed ovipositor (Figs 1C, 8). Reservoir of spermatheca kidney-shaped with short spermathecal duct (Fig. 8). Distribution. Species of Guapinannus are documented from Central and South America, ranging from Veracruz in Mexico to the southern part of Brazil. The greatest species diversity is found in Central America and the northern areas of South America. Specimens have been collected in tropical wet or moist forest, including cloud forest and mixed hardwood forest, from elevation ranging from 5 meters above sea level in Panama to more than 200 meters in Chiapas, Mexico. Collecting method. The majority of specimens for which collection method was recorded were collected using leaf litter sifting and Berlese and Winkler extraction methods. A small number of specimens are derived from Malaise and flight intercept traps. Discussion. The forewing shape, structure with distinct claval suture, and venation set apart Guapinannus from all other genera of Schizopteridae. While Humpatanannus and related genera have not been included in phylogenetic analyses, genitalic features are strikingly different in the two groups, and they are unlikely to be close relatives. The non-genitalic morphology of species of Guapinannus is fairly uniform. Most species are mid-sized and ovoid with some variation, with only few species being distinctly smaller or bigger or markedly elongated ovoid. Some forewing venation features are fairly distinctive and useful as diagnostic characters at the species level, among them the width of the C+Sc and the relative proportion of certain cells. As in many other genera of Schizopteridae, the bulk of diagnostic features is derived from male genitalic structures, in particular the length, curvature, and tip shape of the vesica, shape of the anophoric process, where present, and shape of the pygophore that is expanded into a distinct lobe on the right side in some species. We described as new two species for which only female specimens are known (G. dispar, sp. n. and G. minutus, sp. n.). Both are distinctive based on non-genitalic morphology and G. dispar, sp. n. in addition is geographically isolated from all other known species of Guapinannus. As part of this study we discovered a so far undocumented structure on the forewings that is restricted to males that we interpret as a novel male-specific wing organ. This wing organ is located on the posterior margin of the corium, associated with An1, consist of a strongly inflated semicircular or elongate area and in most species features a median notch. SEM observation of this wing organ in one species (Fig. 6D) shows a tuft of hairlike structures that may represent a trichome. Based on our observations, males of all species of Guapinannus possess this wing organ. Among closely related taxa (based on Knyshov et al., 2020), Caucanannus also have a male-specific wing organ, although in a different position on the forewing (Weirauch et al., 2020), while wing organs have so far not been documented in Luachimonannus and Kokeshia., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on pages 263-265, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718, {"references":["Wygodzinsky, P. (1951) Descripcion de generos y especies nuevos de la familia Cryptostemmatidae (Hemiptera). Revista Brasileira de Biologia, 11, 259 - 270.","Knyshov, A., Weirauch, C. & Hoey-Chamberlain, R. (2020) Phylogenetic relationships and revised classification of the true bug infraorder Dipsocoromorpha (Insecta: Hemiptera: Heteroptera), Cladistics. [early view] https: // doi. org / 10.1111 / cla. 12435","Weirauch, C, Hoey-Chamberlain, R. & Knyshov, A. (2020) Four new genera of Schizopteridae (Hemiptera: Heteroptera) from the Afrotropical and Neotropical regions. Zootaxa, 4768 (1), 95 - 111. https: // doi. org / 10.11646 / zootaxa. 4768.1.6"]}

Guapinannus sinuosus, sp. n. (Figs 3, 7, 9) Holotype. Male. MEXICO: Chiapas: Palenque, 15.63333°N 92.65°W, 1640m, 02 Jul 1983 – 30 Jul 1983, S. B. Peck, J. Kukalova-Peck, R. S. Anderson, 1♂ (UCR _ ENT 00102050) (FMNH). Paratypes: MEXICO: Chiapas: El Bosque (6.6 mi SW), 17.01667°N 92.78333°W, 1463 m, 29 Aug 1973, A. F. Newton, 7♀ (UCR _ ENT 00101241, UCR_ENT 00102051-UCR_ENT 00102053) (FMNH). Playón de la Gloria, 16.16028°N 90.90139°W, 160 m, 26 Jun 2008, collector unknown, 1♂ (UCR _ ENT 00120069) (MTEC). Etymology: Named for the s-shaped vesica after “sinuosus” Latin for bending. Diagnosis: Recognized by the mid-sized body length (1.49 mm), C+Sc margin broad anteriorly and more slender posteriorly, distinctly double s-shaped margin of rc1, rc, tc cells (black arrow) (Fig. 3), and relatively tick and s-shaped vesica (Fig. 7). Description: Male: Total body length 1.49 mm; length from posterior margin of pronotum to posterior wing margin 0.9 mm; greatest width across pronotum 0.54 mm. General coloration: uniformly pale. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum small, evenly distributed; scutellum in dorsal view gradually narrowed to tip, tip pointed. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin rounded, vein-tracing areoles on proximal part of wing absent; C+Sc uniformly broad; distal process of R weakly developed; An1 wing organ on corium strongly inflated, without notch articulating with claval process, rounded, without median notch; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 5 times as long as wide; tc about 3 times as long as wide; distal margin of rc1, rc, and tc cells double s-shaped; rc2–3 almost triangular; cub almost reaching wing margin; dc1 with basal portion relatively wide and short. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, not reaching right pygophore margin, thick at midpoint, tapering to acute tip, close to tip slightly s-shaped; anophoric ridge with process, process short, with single lobe curved, tip pointed; right paramere curved, tapering, basal process broad; left paramere straight, tapering, basal process narrow. Female: As in male and generic description. Collecting method and habitat: Collected using Berlese extraction, flight intercept trap, and sifting leaf litter in closed forest, cloud forest with pines, mature wet forest, and rainforest.

Guapinannus tatumbia, sp. n. (Figs 4, 7, 9) Holotype. Male. HONDURAS: Francisco Morazan: Tatumbia Montaña del Aguacate, 14°N 87.08333°W, 1425m, 14 Mar 1996, R. Cave, 1♂ (UCR _ ENT 00078556) (EAPZ). Paratypes: HONDURAS: Francisco Morazan: Tatumbia Montaña del Aguacate, 14°N 87.08333°W, 1425m, 14 Mar 1996, R. Cave, 5♂ (UCR _ ENT 00078555, UCR_ENT 00078557-UCR_ENT 00078560), 5♀ (UCR _ ENT 00078561 -UCR_ENT 00078565) (EAPZ). Olancho: La Union Parq Nac La Maralla, 15.07949°N 86.76175°W, 1060m, 14 Sep 1994, R. Cordero, 1♀ (UCR _ ENT 00078566) (EAPZ). Etymology: Named for the type locality “Tatumbia Montana” in Honduras (noun in apposition). Diagnosis: Recognized by the mid-sized body length (1.51 mm), elongate oval outline of forewings, inflated tip of scutellum (black arrow) (Fig. 4), vesica relatively thick and straight with acute tip, not reaching pygophore margin, and without anophoric process (Fig. 7). Description: Male: Total body length 1.51 mm; length from posterior margin of pronotum to posterior wing margin 1.27 mm; greatest width across pronotum 0.62 mm; greatest width across forewings 0.88 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long; setae on posterior margin of forewing extremely long. Structure: Head: Eye about one third as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum small, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip rounded. Forewing: With outline broad anteriorly; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing absent; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium inflated, elongate, with median notch, without notch articulating with claval process; rc1 about 3 times as long as wide, much wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2–3 trapezoidal; cub almost reaching wing margin; dc1 with basal portion relatively wide and short. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, not reaching right pygophore margin, thick at midpoint, tapering to acute tip, close to apex fairly straight; anophoric ridge without process; right paramere curved, broad throughout, basal process broad; left paramere straight, tapering, basal process narrow. Female: As in male and generic description. Collecting method and habitat: Unknown.

Guapinannus robustus, sp. n. (Figs 3, 7, 9) Holotype. Male. MEXICO: Chiapas: RSV A El Triunfo, Est. El Triunfo, 16.20286°N 91.86044°W, 1950 m, 08 Aug 1996 – 11 Aug 1996, A. R. Gillogly, 1♂ (UCR _ ENT 00094282) (UCR). Etymology: Named for the extremely thick vesica after “robustus” Latin for robust. Diagnosis: Recognized by the mid-sized body length (1.49 mm), small vein-tracing areoles distributed along scc, rc and rc1 (black arrow) (Fig. 3), slight extensions of the right pygophore margin (black star), and very thick vesica with blunt tip (Fig. 7). Description: Male: Total body length 1.49 mm; length from posterior margin of pronotum to posterior wing margin 1.1 mm; greatest width across pronotum 0.28 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long; setae on posterior margin of forewing extremely long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view gradually narrowed to tip, tip rounded. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin rounded, vein-tracing areoles small, distributed along scc, rc and rc1; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium strongly inflated, triangular, with median notch, without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 5 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 trapezoidal; cub almost reaching wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia (Fig. 7): Pygophore with slightly extended right margin; vesica with about half loop, not reaching right pygophore margin, very thick at midpoint, tapering to blunt tip, close to apex slightly s-shaped; anophoric ridge with process, process short, with single lobe relatively straight, tip pointed; right paramere curved, broad throughout, basal process broad; left paramere straight, tapering, basal process narrow. Female: Unknown. Collecting method: Collected using a flight intercept trap.

Guapinannus plurilobus, sp. n. (Figs 3, 7, 9) Holotype. Male. PERU: Amazonas: Pacaya–Samiria National Reserve, 5.2012°S 74.59319°W, 280m, 07 Nov 1980, CI. Vaucher, 1♂ (UCR _ ENT 00089590) (MHNG). Paratypes: PERU: Amazonas: Pacaya–Samiria National Reserve, 5.2012°S 74.59319°W, 280m, 07 Nov 1980, CI. Vaucher, 4♂ (UCR _ ENT 00089589, UCR_ENT 00089591-UCR_ENT 00089593), 5♀ (UCR _ ENT 00089594 - UCR_ENT 00089598) (MHNG). Unknown locality in Peru, no date provided, collector unknown, 1♀ (UCR _ ENT 00089582) (MHNG). Etymology: Named for the multiple lobes on the anophoric process after “pluri” and “lobus” Latin for multiple lobes. Diagnosis: Recognized by the small body length (1.14 mm), relatively short rc1, rc, and tc cells (black star) (Fig. 3), slender vesica with one loop, and multilobed anophoric process, with combination of blunt and acute tips (black arrow) (Fig. 7). Description: Male: Total body length 1.14 mm; length from posterior margin of pronotum to posterior wing margin 0.8 mm; greatest width across pronotum 0.44 mm; greatest width across forewings 0.67 mm. General coloration: uniformly pale. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip rounded. Forewing: With outline broad anteriorly; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing absent; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium weakly inflated, elongate, with median notch; without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc slightly wider than other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 almost triangular; cub almost reaching wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with one loop, not reaching right pygophore margin, relatively slender at midpoint, tapering to blunt tip, close to apex rounded; anophoric ridge with process, process long, with multiple lobes, tip blunt; right paramere curved, broad throughout, basal process broad; left paramere straight, tapering, basal process narrow. Female: As in male and generic description. Collecting method: Collected using Winkler extraction.

Guapinannus falcis, sp. n. (Figs 2, 7, 8, 9) Holotype. Male. COSTA RICA: Heredia: 6 km ENE Vara Blanca, 10.18333°N 84.11667°W, 2000 m, 16 Mar 2002, INBio-OET-ALAS, 1♂ (UCR _ ENT 00014616) (INBIO). Paratypes: COSTA RICA: Alajuela: Upala Co.: Bijagua, San Miguel, Finca Inti-Aura, 10.75778°N 85.00939°W, 300 m, 16 Apr 2011 – 20 May 2011, I. Chacon, 1♂ (UCR _ ENT 00014608) (INBIO). Heredia: Sarapiqui Co.: P.N.B. Carrillo 5 km. E Vara Blanca, 10.17226°N 84.11213°W, 2100 m, 20 Apr 2002, INBio-OET-ALAS, 1♀ (UCR _ ENT 00014611) (INBIO). P.N.B. Carrillo 6 km. ENE Vera Blanca, 10.15°N 83.91667°W, 2000 m, 17 Feb 2002 – 21 Feb 2002, INBio-OET-ALAS, 1♀ (UCR _ ENT 00014612) (INBIO); 10 Mar 2002, INBio-OET-ALAS, 1♀ (UCR _ ENT 00014610), 1♂ (UCR _ ENT 00014613) (INBIO). 6 km ENE Vara Blanca, 10.18333°N 84.11667°W, 2000 m, 16 Mar 2002, INBio-OET-ALAS, 1♀ (UCR _ ENT 00014614) (INBIO); 22 Mar 2002, INBio-OET-ALAS, 4♀ (UCR _ ENT 00014618 -UCR_ENT 00014621) (INBIO). Etymology: Named for sickle shaped vesica after “flax” Latin for sickle. Diagnosis: Recognized by the mid-sized body length (1.56 mm), dense vestiture, part of An1 and Cu forming posterodistal margin of tc more pronounced than other veins (black arrow) (Fig. 2), long, relatively slender vesica, with s-shaped bent proximal of acute tip, and short, curved anophoric process with pointed tip (black arrow) (Fig. 7). Most similar to G. bierigi, sp. n., that is also known from Costa Rica, but distinguished by the much smaller distance of cub from the wing margin compared to G. bierigi, sp. n. Description: Male: Total body length 1.56 mm; length from posterior margin of pronotum to posterior wing margin 1.27 mm; greatest width across pronotum 0.65 mm; greatest width across forewings 0.87 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense, long setae; setae on posterior margin of forewing long. Structure: Head: Eye about one third as high as head; muscle scars not observed. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum large, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip pointed. Forewing: With outline broad anteriorly, elongate oval; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing absent; C+Sc uniformly broad; distal process of R pronounced; An1 wing organ on corium inflated, elongate, with median notch; without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 5 times as long as wide; tc more than 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc more pronounced than other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 almost triangular; cub almost reaching wing margin; dc1 with basal portion relatively wide and short. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, surpassing right pygophore margin, relatively slender at midpoint, tapering to acute tip, close to apex slightly s-shaped; anophoric ridge with process, process short, with single lobe curved, tip pointed; right paramere curved, broad throughout, basal process broad; left paramere straight, tapering, basal process broad. Female: As in male and generic description. Collecting method and habitat: Collected using Malaise trap in unknown habitat., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on page 272, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718

Guapinannus graziae, sp. n. (Figs 3, 7, 9) Holotype. Male. NICARAGUA: Region Autonoma del Atlantico Sur: 13km WNW Rama, 12.19472°N 84.33667°W, 190 m, 18 Apr 2011, J. T. Longino, 1♂ (UCR _ ENT 00102045) (FMNH). Paratypes: NICARAGUA: Region Autonoma del Atlantico Sur: 13km WNW Rama, 12.19472°N 84.33667°W, 190 m, 18 Apr 2011, J. T. Longino, 3♀ (UCR _ENT 00102046-UCR_ENT 00102048), 1♂ (UCR _ENT 00101103) associated with Carduus occidentalis Nutt. (Asteraceae), 1♂ (UCR _ENT 00102049) (FMNH). Rivas: Matagalpa: RN Cerro Musún, 12.95944°N 85.225°W, 800m, 01 May 2011, collector unknown, 1♂ (UCR _ENT 00101361) (FMNH). Etymology: Named after our dear mentor, colleague, and friend, the grande dame of South American heteropteran systematics, Jocelia Grazia. Diagnosis: Recognized by the small body length (1.16 mm), very broad C+Sc margin (white arrow) (Fig. 3), slightly s-shaped vesica, and short, curved anophoric process (black arrow) (Fig. 7). Description: Male: Total body length 1.16 mm; length from posterior margin of pronotum to posterior wing margin 0.89 mm; greatest width across pronotum 0.56 mm; greatest width across forewings 0.69 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip pointed. Forewing: With outline broad anteriorly; membrane contributing about one third of forewing length, posterior distal margin squarish, veintracing areoles on proximal part of wing absent; C+Sc very broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium strongly inflated, elongate, with median notch, without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells s-shaped; rc2–3 trapezoidal; cub almost reaching wing margin; dc1 with basal portion relatively wide and short. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, not reaching right pygophore margin, thick at midpoint, tapering to acute tip, close to apex slightly s-shaped; anophoric ridge with process, process short, with single lobe curved, tip pointed; right paramere curved, tapering, basal process narrow; left paramere straight, tapering, basal process narrow. Female: As in male and generic description. Collecting method and habitat: Collected using Winkler extraction in second growth forest and tropical wet forest., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on pages 272-273, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718

Guapinannus dispar, sp. n. (Figs 2, 9) Holotype. Female. BRAZIL: Mato Grosso: Jacare Alto Xingu, 10.54341°S 53.75637°W, 330m, 01 Nov 1965, Alvarenga & Werner, 1♀ (UCR _ ENT 00090847) (FMNH). Etymology: Named for the characteristic wing venation after “dispar” Latin for different. Diagnosis: Recognized by the very small size (total body length 0.66 mm), forewing with distinct honey-comb pattern (black arrow), absence of An1/2 claval process, expanded scc divided by sinuous, transverse vein (white arrow), length and shape of rc1 (black star), distinctly surpassing apex of rc and tc, rectangular rc2–3 (white star), and slender and elongate cub (grey star) and tc (Fig. 2). Description: Female: Total body length 0.66 mm; length from posterior margin of pronotum to posterior wing margin 0.51 mm; greatest width across pronotum 0.28 mm; greatest width across forewings 0.42 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with sparse, short setae; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars small. Thorax: Anterior pronotal margin concave, posterior margin almost straight; pits on pronotum large, evenly distributed; scutellum in dorsal view gradually narrowed to tip, tip pointed. Forewing: with outline broad anteriorly, elongate oval; with distinct honey-comb pattern; membrane contributing less than one third of forewing length, posterior distal margin rounded, vein-tracing areoles on proximal part of wing absent; C+Sc narrow, wider at widest part of wing; distal process of R weakly developed; An1/2 process on clavus absent; An1 wing organ absent (as in all females); rc1 about 4 times as long as wide, much wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells angular s-shaped; rc2-3 rectangular; cub removed from wing margin; dc1 with basal portion relatively wide and short. Legs: Claws short. Genitalia: As in generic description. Male: Unknown. Collecting method and habitat: Unknown. Discussion: This species is set apart from all other species of Guapinannus both in terms of morphology (size, forewing sculpture, wing venation) and distribution (the only documented species of Guapinannus from Brazil). We decided to include this species into Guapinannus and to somewhat expand the concept of this genus, because G. dispar, sp. n., is more similar to species of “core” Guapinannus than to any other species of Schizopteridae we have seen from the Neotropical region. Once male specimens become available for this species, our hypothesis should be further tested., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on pages 270-272, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718

Guapinannus castigatus, sp. n. (Figs 2, 7, 9) Holotype. Male. MEXICO: Chiapas: Palenque, 15.63333°N 92.65°W, 2500m, 02 Jul 1983 – 30 Jul 1983, S. B. Peck, J. Kukalova-Peck, R. S. Anderson, 1♂ (UCR _ ENT 00101251) (FMNH). Etymology: Named for the stout vesica after “castigatus” Latin for stout. Diagnosis: Recognized by the mid-sized body length (1.48 mm), elongate oval forewing outline, wide C+Sc (black arrow), small vein-tracing areoles along scc, rc, rc1 (white arrow) (Fig. 2), slightly extended right pygophore margin (black star), vesica relatively short and thick, tapering into acute tip, and without anophoric process (Fig. 7). Similar to G. auriculus, sp. n. in length and shape of vesica but differentiated by differentiated from that species by the slender and long rc1. Description: Male: Total body length 1.48 mm; length from posterior margin of pronotum to posterior wing margin 1.27 mm; greatest width across pronotum 0.66 mm; greatest width across forewings 0.88 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view gradually narrowed to tip, tip pointed. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing present; areoles small, distributed along scc, rc, rc1, and tc; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium strongly inflated, elongate, with median notch, without notch articulating with claval process; scc about 2 times as wide as long; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 trapezoidal; cub almost reaching wing margin; dc1 with basal portion relatively wide and short. Legs: Claws long. Genitalia (Fig. 7): Pygophore with slightly extended right margin; vesica with about half loop, not reaching right pygophore margin, thick at midpoint, tapering to acute tip, close to apex rounded; anophoric ridge without process; right paramere curved, tapering, basal process narrow; left paramere straight, tapering, basal process narrow. Female: Unknown. Collecting method and habitat: Collected using flight intercept trap in rainforest.

Guapinannus minutus, sp. n. (Figs 3, 9) Holotype. Female. COSTA RICA: Heredia: Sarapiqui Co.: P.N. Braulio Carrillo, 16 Km SSE La Virgen, 10.26784°N 84.084°W, 1050 m, 20 Feb 2001 – 23 Feb 2001, INBio-OET-ALAS, 1♀ (UCR _ ENT 00014609) (INBIO). Etymology: Named for the small body size after “minutus” Latin for small. Diagnosis: Recognized by the small size (1.14 mm), inflated tip of scutellum (white arrow), well-defined proximal part of Cu (black arrow), and dark coloration (female-based) (Fig. 3). Description: Female: Total body length 1.14 mm; length from posterior margin of pronotum to posterior wing margin 0.90 mm; greatest width across pronotum 0.52 mm; greatest width across forewings 0.71 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip inflated. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin rounded, vein-tracing areoles on proximal part of wing absent; C+Sc uniformly fairly broad; distal process of R pronounced; An1 wing organ on corium absent (as in all females); rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 almost triangular; cub almost reaching wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia: As in generic description. Male: Unknown. Collecting method and habitat: Unknown.

Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae) was described based on a single female specimen from Costa Rica. Some additional specimens representing this genus have since become available and were incorporated into a comparative male genitalic study across Dipsocoromorpha and into combined molecular and morphological hypotheses of the infraorder. However, the species-level diversity of Guapinannus has remained unexplored and undocumented. Based on examination of 264 specimens from central and South America, we here revise the taxonomy of Guapinannus, describing 19 species as new (Guapinannus anaticulus, sp. n.; Guapinannus artus, sp. n.; Guapinannus auriculus, sp. n.; Guapinannus castigatus, sp. n.; Guapinannus clava, sp. n.; Guapinannus dispar, sp. n.; Guapinannus falcis, sp. n.; Guapinannus graziae, sp. n.; Guapinannus minutus, sp. n.; Guapinannus orbiculatus, sp. n.; Guapinannus plurilobus, sp. n.; Guapinannus policis, sp. n.; Guapinannus robustus, sp. n.; Guapinannus sinuosus, sp. n.; Guapinannus tatumbia, sp. n.; Guapinannus tenuis, sp. n.; Guapinannus tergus, sp. n.; Guapinannus trilobus, sp. n.; Guapinannus uncus, sp. n.). In addition, we provide photos of the female holotype of Guapinannus bierigi Wygodzinsky, 1951, SEM documentation for Guapinannus clava, sp. n., habitus photos and a map for all species, and line drawings of male genitalic features for all species for which males are known.

Guapinannus orbiculatus, sp. n. (Figs 3, 7, 9) Holotype. Male. PANAMA: Bocas del Toro: Bocas del Toro: Almirante, trail to dam on Nigua Creek, 9.29792°N 82.4211°W, 32m, 31 Mar 1959, H. S. Dybas, 1♂ (UCR _ ENT 00090718) (FMNH). Paratypes: PANAMA: Bocas del Toro: Bocas del Toro: Almirante, trail to dam on Nigua Creek, 9.29792°N 82.4211°W, 12m, 31 Mar 1959, H. S. Dybas, 1♂ (UCR _ ENT 00097813), 4♀ (UCR _ ENT 00097814 -UCR_ENT 00097817), 2 nymphs (UCR _ ENT 00097818, UCR_ENT 00097819) (FMNH). Chiriqui: Near Nueva California W. of Finca Palo Santo, 8.5°N 82.41667°W, 1524 m, 10 Mar 1959, H. S. Dybas, 1♂ (UCR _ ENT 00090796) (FMNH). Diagnosis: Recognized by the small mid-sized body length (1.26 mm), elongate body shape (Fig. 3), one-loop, relatively thick vesica with pointed tip, and short, slender, pointed anophoric process (black arrow) (Fig. 7). Description: Male: Total body length 1.26 mm; length from posterior margin of pronotum to posterior wing margin 0.9 mm; greatest width across pronotum 0.53 mm; greatest width across forewings 0.68 mm. General coloration: uniformly pale. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye about one third as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin slightly concave; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip rounded. Forewing: With outline elongate oval; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing absent; C+Sc uniformly fairly broad; distal process of R pronounced; An1 wing organ on corium inflated, elongate, with median notch; without notch articulating with claval process; rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc slightly wider than other veins; distal margin of rc1, rc, and tc cells double s-shaped; rc2-3 trapezoidal; cub almost reaching wing margin; dc1 horse head-shaped. Legs: Claws long. Genitalia (Fig. 7): Pygophore without lobe on right side; vesica with between half and one loop, not reaching right pygophore margin, thick at midpoint, tapering to acute tip, close to apex rounded; anophoric ridge with process, process short, slender, with single lobe curved, tip pointed; right paramere curved, tapering, basal process broad; left paramere straight, tapering, basal process narrow. Female: As in male and generic description. Collecting method and microhabitat: Collected using Berlese and Winkler extraction from the cortex of a hollowed palm stump and from floor litter at base of a log and cut stump. Etymology: Named for the rounded vesica after “orbiculatus” Latin for round., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on page 274, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718

Guapinannus bierigi Wygodzinsky, 1951 (Figs 1, 9) Diagnosis: Recognized by the mid-sized body length (1.45 mm), part of An1 and Cu forming posterodistal margin of tc more pronounced than other veins, and the great distance of cub from the margin of the wing. Most similar to G. falcis, sp. n., that is also known from Costa Rica, but distinguished by the much greater distance of cub from the wing margin compared to G. falcis, sp. n. Re-description: Female: Total body length 1.45 mm, greatest width across pronotum 0.67 mm. General coloration: mix of pale and darker brown or reddish brown. Vestiture: Dorsum with sparse setae, setae short; setae on posterior margin of forewing long. Structure: Head: Eye almost half as high as head; muscle scars large. Thorax: Anterior pronotal margin concave, posterior margin straight; pits on pronotum medium-sized, evenly distributed; scutellum in dorsal view abruptly narrowed to tip, tip pointed. Forewing: With outline broad anteriorly; membrane contributing about one third of forewing length, posterior distal margin squarish, vein-tracing areoles on proximal part of wing absent; C+Sc broad, wider at widest part of wing; distal process of R pronounced; An1 wing organ on corium absent (as in all females); rc1 about 3 times as long as wide, slightly wider anteriorly; rc about 4 times as long as wide; tc about three times as long as wide; part of An1 and Cu forming posterodistal margin of tc more pronounced than other veins; distal margin of rc1, rc, and tc cells s-shaped; rc2-3 almost triangular; cub removed from wing margin; dc1 with basal portion fairly narrow and elongate. Legs: Claws long. Genitalia: As in generic description (Fig. 1). Male: Unknown. Specimens examined: Holotype: COSTA RICA: Limon: Guapiles, 10.2167°N 83.7833°W, 268 m, 05 Feb 1946 – 13 Feb 1946, A. Bierig, 1♀ (AMNH _ IZC 00150354) (AMNH)., Published as part of Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, pp. 261-286 in Zootaxa 4958 (1) on pages 268-269, DOI: 10.11646/zootaxa.4958.1.14, http://zenodo.org/record/4691718, {"references":["Wygodzinsky, P. (1951) Descripcion de generos y especies nuevos de la familia Cryptostemmatidae (Hemiptera). Revista Brasileira de Biologia, 11, 259 - 270."]}

Although they are a valuable source of specimens, insect natural history collections continue to be under‐utilized in molecular systematics, mostly due to difficulties in obtaining DNA sequences. Old specimens or specimens stored under suboptimal conditions are intractable for traditional Sanger sequencing. In this study we use an inexpensive hybrid capture with in‐house generated baits to retrieve commonly utilized ribosomal and mitochondrial loci from old museum specimens and combine them with a Sanger‐generated dataset comprising recently collected material. We focus on the Corixidea genus group (Schizopteridae), which comprises rarely collected, small (1–2 mm) and primarily tropical insects of which only c. 10–20% of the species have been described. A molecular phylogeny is needed to resolve relationships and revise the genus‐level classification to correctly place the c. 150 yet to be described species. Applying this approach, we constructed a dataset, containing 101 taxa, 11 of which were preserved in low‐percentage ethanol, 48 are dry and point‐mounted, and 40 are > 20 years old at DNA extraction. The obtained data proved sufficient for reconstructing a well‐supported phylogeny with c. 50% of the predicted diversity, and for the oldest successfully sequenced specimen (95 years) to be unambiguously placed in that phylogeny. We confirmed monophyly of the Corixidea genus group, showed paraphyly of the genus Corixidea, and recovered nine well‐supported clades within the group. Ancestral character states of selected morphological features were inferred and used to re‐examine primary homology hypotheses and inform an upcoming taxonomic revision.

Gathering genetic data for rare species is one of the biggest remaining obstacles in modern phylogenetics, particularly for megadiverse groups such as arthropods. Next generation sequencing techniques allow for sequencing of short DNA fragments contained in preserved specimens >20 years old, but approaches such as whole genome sequencing are often too expensive for projects including many taxa. Several methods of reduced representation sequencing have been proposed that lower the cost of sequencing per specimen, but many remain costly because they involve synthesizing nucleotide probes and target hundreds of loci. These datasets are also frequently unique for each project and thus generally incompatible with other similar datasets. Here, we explore utilization of in-house generated DNA baits to capture commonly utilized mitochondrial and ribosomal DNA loci from insect museum specimens of various age and preservation types without the a priori need to know the sequence of the target loci. Both within species and cross-species capture are explored, on preserved specimens ranging in age from one to 54 years old. We found most samples produced sufficient amounts of data to assemble the nuclear ribosomal rRNA genes and near complete mitochondrial genomes and produce well-resolved phylogenies in line with expected results. The dataset obtained can be straightforwardly combined with the large cache of existing Sanger-sequencing-generated data built up over the past 30 years and targeted loci can be easily modified to those commonly used in different taxa. Furthermore, the protocol we describe allows for inexpensive data generation (as low as ~$35/sample), of at least 20 kilobases per specimen, for specimens at least as old as ~1965, and can be easily conducted in most laboratories. If widely applied, this technique will accelerate the accurate resolution of the Tree of Life especially on non-model organisms with limited existing genomic resources.

Because species diversity of the small true bug family Schizopteridae is greatest in tropical and subtropical areas, it is not surprising that only four species have been described from the United States. As part of a larger project on the taxonomy and phylogenetics of Schizopteridae, 178 specimens from the United States were examined. This material contained representatives of the previously described species Glyptocombussaltator Heidemann, 1906, Corixideamajor McAtee & Malloch, 1925, Nannocorisarenarius Blatchley, 1926, and Schizopterabispina McAtee & Malloch, 1925, but also six undescribed species. These new taxa are described as Glyptocombushalbertaesp. n., Glyptocombussuterisp. n., Nannocorisanophorussp. n., Nannocorisbrevipilussp. n., Schizoptera (Cantharocoris) rileyisp. n., and Schizoptera (Schizoptera) henryisp. n. Habitus images and genitalic illustrations of the previously described and the new species are provided as well as a map showing distribution ranges of these species in the United States and Mexico. To provide a comprehensive treatment of the small genus Glyptocombus Heidemann, 1906, Glyptocombusmexicanussp. n. is also described that, to our knowledge, occurs only in Mexico, and the female of one additional undescribed Glyptocombus species is documented from Mexico.

Automated insect identification systems have been explored for more than two decades but have only recently started to take advantage of powerful and versatile convolutional neural networks (CNNs). While typical CNN applications still require large training image datasets with hundreds of images per taxon, pretrained CNNs recently have been shown to be highly accurate, while being trained on much smaller datasets. We here evaluate the performance of CNN-based machine learning approaches in identifying three curated species-level dorsal habitus datasets for Miridae, the plant bugs. Miridae are of economic importance, but species-level identifications are challenging and typically rely on information other than dorsal habitus (e.g., host plants, locality, genitalic structures). Each dataset contained 2–6 species and 126–246 images in total, with a mean of only 32 images per species for the most difficult dataset. We find that closely related species of plant bugs can be identified with 80–90% accuracy based on their dorsal habitus alone. The pretrained CNN performed 10–20% better than a taxon expert who had access to the same dorsal habitus images. We find that feature extraction protocols (selection and combination of blocks of CNN layers) impact identification accuracy much more than the classifying mechanism (support vector machine and deep neural network classifiers). While our network has much lower accuracy on photographs of live insects (62%), overall results confirm that a pretrained CNN can be straightforwardly adapted to collection-based images for a new taxonomic group and successfully extract relevant features to classify insect species.

Perittonannus gen. n. urn:lsid:zoobank.org:act: 207E33A1-959D-4B09-9909-C29FDC791878 Figures 1, 2D, E, 3 G���I, 6B, 7A Type species. Perittonannus antiquus sp. n. Diagnosis. Distinguished among Schizopteridae by the relatively small body size (0.96-1.13 mm), small round- ed head about as long as high in lateral view, medium-sized eye in lateroventral position, slender four-segmented labium (Fig. 1), R with three branches, subcostal vein, R1 and R2 fused just beyond midpoint of forewing, rc3 boomerang-shaped, dc1 large, rc and tc cells of similar subrectangular shape and size, tc with four emanating veins (Fig. 2D), long, robust, and very setose legs (Fig. 1), tarsal formula 2-2-2 (Fig. 3 G-I), symmetrical abdomen with dorsum and ventrum equally sclerotized, genitalia sinistrally directed, laterotergites 9 present on both sides as free-moving appendages with enlarged apices, short thick vesica with apical funnel-shaped expansion, conjunctiva with heavily sclerotized quadrate sclerite on left side of pygophore (see Knyshov et al. [2018] their Figs 1o, 4d, 14b), and the well-developed ovipositor. Description. Male: macropterous, suboval, relatively small body size (0.96-1.13 mm). Coloration (Fig. 1). Body dark brown, antenna pale yellowish, labium pale yellowish with dark brown apex, forewing dark brown with broad white transverse band across proximal portion of subcostal cell, basal cell, and distal half of clavus, legs pale yellowish. Vestiture (Fig. 1). Body with dense cover of microtrichia, some longer semierect setae lateroventrally on head and laterally on thorax, and medium-length subadpressed setae dorsally on head and pronotum, legs densely beset with long, fairly stout, semirect setae. Structure. Head (Fig. 1): small and rounded in dorsal view, about as long as high in lateral view, synthlipsis about 5.5 times as wide as eye, eye in lateroventral position, largely obscured in dorsal view, prominent in ventral view; labium four-segmented, reaching apex of middle coxa, first segment inflated, second to fourth segments very slender and of similar diameter; antenna with short scape and pedicel (flagellomeres missing). Thorax (Fig. 1): pronotum broad and short, slightly less than half as long as wide, collar narrow, anterior and posterior pronotal lobes not demarcated, posterior margin in dorsal view almost straight, scutellum small and narrow, with tapering apex, metathorax without obvious gland evaporatory structures. Forewing (Fig. 2D): macropterous, with slender scc, R with three branches, Sc, R1 and R2 fused just beyond midpoint of forewing, rc3 narrow and boomerang-shaped, dc1 large, rc and tc of similar subrectangular shape and size, cub oval, relatively small; Sc and R1 heavily sclerotized to point of fusion with R2, Sc distinct to tip of forewing and Cu. Legs: long and robust (Fig. 1), subapical combs on fore-, middle, and hind tibiae (Fig. 3 G-I), hind tibia with row of slender capitate setae (Fig. 3H, I); tarsal formula 2-2-2 (Fig. 3 G-I), fore- and middle tarsus as slender as hind tarsus incrassate (Fig. 3 G-I), first tarsomeres very short, second ~3-4 times longer than first, claws slender and long, ~1/3 of second tarsomere length; bladderlike arolium absent, parempodia with tapering apex. Abdomen and genitalia: Documented and described in Knyshov et al. (2018) based on specimen UCR_ENT 00014811 (their Figs 1o, 4d, 14b, and 19e): ���Pregenital abdomen. Symmetrical, not laterally curved, dorsum, and ventrum equally sclerotized. All abdominal segments equally sclerotized. Dorsal surface smooth. Pregenital laterotergites indistinct. Two pairs of spiracles present (segments 7���8), located on lateral margins of sternum (segment 7), or on lateral margins of mediotergite (segment 8). Scars of DAG orifices present medially along anterior margin of mediotergite 7. Pregenital abdomen modifications. Mediotergite 7 symmetrical, short. Laterotergites indistinct. Sternum 7 symmetrical, long. Genitalia. Sinistrally directed. Mediotergite 8 symmetrical, short. Laterotergites 8 indistinct. Sternum 8 indistinct. Pygophore symmetrical, oval, open dorsally, mediotergite 9 indistinct. Laterotergites 9 distinct, nearly symmetrical, elongated. Parameres asymmetrical, left larger than right. Left paramere elongated, basal process elongated and rounded, apical process elongated and rounded. Right paramere elongated, basal process short and rounded, apical process elongated and rounded. Basal plates weakly asymmetrical. Phallosoma reduced. Conjunctiva heavily sclerotized, represented by large quadrate sclerite and one short rounded appendage. Vesica with narrow base and coiled thick short distal region, expanding around apex into funnel. Anophore symmetrical, without processes. Anal tube membranous.��� Female (Fig. 1). Similar to male in coloration and vestiture, but submacropterous (Fig. 2E) and distinctly smaller than male (0.68���0.86 mm) (Tab. 1); tarsal formula 2-2-2. Genitalia (Fig. 6B): well-developed, sclerotized ovipositor formed by interlocked gonapophyses 8 and 9, and styloids; gonapophysis 8 slightly longer than gonapophysis 9, both with sharp apex and without teeth or large setae; styloids represented by six interconnected sclerites median to gonapophysis 8; spermathecal gland small and spherical; spermathecal gland duct straight; spermathecal reservoir large and spherical; spermathecal duct relatively short and very narrow, opening into large spheroid membranous pouch. Etymology. From peritt��s (Greek) meaning ���odd��� to reflect the singular morphology of this phylogenetically isolated taxon and ���nannus��� after the Greek king of the same name (Nannus) and resembling n��nos meaning ���dwarf��� (Greek). The gender is masculine. Distribution. Known from Costa Rica. Notes. The isolated phylogenetic position of Perittonannus gen. n. in the combined molecular and morphological analyses by Knyshov et al. (under revision) are reflected in the singular morphology of this taxon. Perittonannus gen. n. is unique among non-hypselosomatine Schizopteridae in the sinistrally oriented male genitalia, among other genitalic features, and by numerous aspects of the forewing venation. A well-developed ovipositor is rare nonhypselosomatine Schizopteridae and in this case is likely plesiomorphic. Perittonannus antiquus sp. n. urn:lsid:zoobank.org:act: 3CAA6CD1-EDDF-4A1A-B31A-BA5D53C1144A Figures 1, 2D, E, 3 G���I, 6B, 7A Diagnosis. As in generic diagnosis. Description. As in generic description. Measurements. See Table 1. Etymology. According to our phylogenetic analyses, this genus and species represent the earliest diverging lineage of non-hypselosomatine Schizopteridae. The species epithet antiquus meaning ���ancient��� was chosen to reflect this phylogenetic position. Specimens examined. Holotype: COSTA RICA: Heredia: 16 km SSE La Virgen, 10.26666 �� N 84.08333 �� W, 1150 m, 09 Mar 2001 ��� 14 Mar 2001, E. G. Riley, 1;m (UCR _ ENT 00093427) (TAMU). Paratypes: COSTA RICA: Heredia: Sarapiqui Co.: Heredia: La Selva Biol. Station, 10.43 �� N 84.02 �� W, 16 Feb 2002 ��� 17 Feb 2002, B. Brown, L. Gonzales, and K. Walker, 1;m (UCR _ ENT 00022675) (LACM). P.N. Braulio Carrillo, 16 Km SSE La Virgen, 10.26784 �� N 84.084 �� W, 1050 m, 20 Feb 2001 ��� 23 Feb 2001, INBio-OET-ALAS, 2;f (UCR _ ENT 00014571, UCR_ENT 00014572) (INBIO); 23 Feb 2001, INBio-OET-ALAS, 3;f (UCR _ ENT 00014574, UCR_ENT 00014577, UCR_ENT 00014579) (INBIO); 14 Mar 2001 ��� 17 Mar 2001, INBio-OET-ALAS, 1;m (UCR _ ENT 00014814) (INBIO); 17 Mar 2001, INBio-OET-ALAS, 1;m (UCR _ ENT 00014812) (INBIO); 17 Mar 2001, INBio-OET-ALAS, 3;f (UCR _ ENT 00014573, UCR_ENT 00014576, UCR_ENT 00014578) (IN- BIO); 19 Mar 2001 ��� 22 Mar 2001, INBio-OET-ALAS, 1;f (UCR _ ENT 00014570) (INBIO). La Selva, 10.42375 �� N 84.02175 �� W, 15 Feb 2002 ��� 16 Feb 2002, Brown, 2;m (UCR _ ENT 00039260) (LACM). La Selva Biological Station, 10.43333 �� N 84.01667 �� W, 16 May 1992, ALAS, Light Trap, 1;m (UCR _ ENT 00109111) (UCR). Limon: Hamburg Farm, 10.25 �� N 83.45 �� W, Nov 1927, Unknown, 1;m (UCR _ ENT 00096869) (AMNH). Rio Sardinas, R. N.F.S. Barra del Colorado, A.C. Tortuguero, 10.69184 �� N 83.71991 �� W, 50 m, Mar 1994, F. Araya, 2;m (UCR _ ENT 00014810, UCR_ENT 00014809) (INBIO). Unknown: Costa Rica, 9.74277 �� N 83.75944 �� W, 1167 m, no date provided, Unknown, 1;f (UCR _ ENT 00014575), 2;m (UCR _ ENT 00014811, UCR_ENT 00014813) (INBIO). Collecting method and habitat. Specimens were collected using Yellow Pan Traps and Malaise Traps in Isthmian-Atlantic moist forest. One specimen was recorded from ���successional plots��� at La Selva Biological Station, suggesting that specimens may also be found in somewhat disturbed habitats. Rimanannus gen. n. urn:lsid:zoobank.org:act: FC5E7FA1-1477-4663-B66B-450DF7FEF4D1 Figures 1���3, 5C, 7B Type species. Rimanannus camerunensis sp. n. Diagnosis. Distinguished among Schizopteridae by the small size (~ 1 mm), medium-sized eyes, four-segment- ed labium (Fig. 1), distinctive wing feature comprising strongly inflated mid-portion of R and M with slit-like rc giving the appearance of a slender coffee bean (Fig. 2F), long costal fracture reaching to apex of bc, tc with five emanating veins, tarsal formula 2-2-3 (Fig. 3 J-L), male sternum 7 shaped into large subgenital plate, laterotergites 9 present on both sides as free-moving appendages with curved apices, aedeagus with one large heavily sclerotized conjunctival appendage, and vesica short and slender (Fig. 5B). Description. Male: macropterous, suboval, small (~ 1 mm). Coloration (Fig. 1). Body and wings uniformly brown, antenna, labium, and legs pale yellow. Vestiture (Fig. 1). Body and forewing smooth, with relatively few medium-length setae, legs moderately setose. Structure. Head (Fig. 1): short and strongly declivous, synthlipsis ~3 times as wide as eye, eye medium-sized; labium four-segmented, reaching base of middle coxa, first segment inflat- ed, second to fourth moderately stout; antenna with short scape and pedicel (flagellomeres missing). Thorax (Fig. 1): pronotum very broad and short, about a quarter as long as wide, collar distinct, longer medially than laterally, anterior and posterior pronotal lobes not demarcated, posterior margin in dorsal view slightly concave, scutellum with spine-like apex, metathorax without obvious gland evaporatory structures. Forewing (Fig. 2F): macropterous, with fairly broad cc, R with 3 branches, R1 forming proximal margin of costal fracture, distinctive wing feature comprising strongly inflated mid-portion of R and M with slit-like rc giving the appearance of a slender coffee bean (Fig. 2F), long costal fracture reaching to apex of bc, bc slender and elongate, tc suboval with five emanating veins, dc elongate and narrow, cub open distally (An1 not reaching Cu); veins surrounding sc, rc, rc1, and rc2 heavily sclerotized (Fig. 2F). Legs: slender (Fig. 1), fore tibia only slightly expanded at apex 9 Fig. 3J), all tibiae subapically with combs (Fig. 3 J-L), hind tibia with row of short, stout, subadpressed setae and with row of erect, more distantly spaced slender setae with capitate apex (Fig. 3L); tarsal formula 2-2-3 (Fig. 3 J-L), with fore- and middle tarsus slightly incrassate (Fig. 3J, K), hind tarsus more slender (Fig. 3L), first tarsomeres very short, second about five times longer than first on fore- and middle leg, second and third hind leg tarsomeres of similar length; fore- and middle tarsi without bladderlike arolium (Fig. 3J, EK), parempodia with tapering apex (Fig. 3L). Abdomen and genitalia (Fig. 5C): pregenital abdomen fairly symmetrical; spiracles located on sterna 6 and 7, and on mediotergite 8; DAG scars minute, located on sides of anterior margin of mediotergite 7 and obscured by mediotergite 6; sternum 7 shaped into subgenital plate, completely covering pygophore from beneath; mediotergite 8 without processes; laterotergites 9 present on both sides as free-moving appendages with curved apices; aedeagus with complex middle portion with tentatively one large heavily sclerotized conjunctival appendage; vesica relatively short and slender, with minute serration near apex; right paramere elongate, L-shaped, with rounded apex; left paramere as long as wide, with rounded apex; anophore sclerotized, devoid of processes. Female. Unknown. Etymology. ���Rima��� from rima (Latin) meaning ���crack, crevice��� for the distinctively inflated R and M portion surrounding the slit-like rc and and ���nannus��� after the Greek king of the same name (Nannus) and resembling n��nos meaning ���dwarf��� (Greek). The gender is masculine. Distribution. Known from two sites in the western part of Cameroon, Mt. Cameroon and Korup National Park. Notes. See under Kamakonocoris gen. n. for discussion of phylogenetic relationship. Rimanannus camerunensis sp. n. urn:lsid:zoobank.org:act: 1D8BCF9E-76A3-475C-895B-6EC1AA79B060 Figures 1, 2F, 3 J���L, 5C, 7B Diagnosis. As in generic diagnosis. Description. As in generic description. Measurements. See Table 1. Etymology. This species is named for its country of origin. Specimens examined. Holotype: CAMEROON: Southwest: Korup National Park, nature trail, 5.00833 �� N 8.86944 �� E, 93 m, 30 Aug 2013 ��� 31 Aug 2013, Forthman, Gordon, Heraty & Weirauch, 1;m (UCR _ ENT 00088644) (UCR). Paratypes: CAMEROON: Southwest: Korup National Park, nature trail, 5.00833 �� N 8.86944 �� E, 93 m, 28 Aug 2013 ��� 30 Aug 2013, Forthman, Gordon, Heraty & Weirauch, 2;m (UCR_ENT 00088115, UCR_ENT 00088116) (UCR). Mt. Cameroon, 4.1175 �� N 9.0716 �� E, 1079 m, 28 Dec 2015, V. Grebennikov, 2;m (UCR_ENT 00127864, UCR_ENT 00127865) (CNC). ~ 8 km N of Mundemba, toward Toko, 4.99994 �� N 8.96401 �� E, 03 Sep 2013, Forthman, Gordon, Heraty & Weirauch, 1;m (UCR_ENT 00087547;ED_2203) (UCR). Collecting method and habitat. Specimens were collected using Yellow Pan Traps in Cross-Sanaga-Bioko coastal forests, a tropical moist broadleaf forest ecoregion of west-central Africa, and from sifting leaf litter in Mount Cameroon and Bioko montane forests., Published as part of Weirauch, Christiane, Knyshov, Alexander & Hoey-Chamberlain, Rochelle, 2020, Four new genera of Schizopteridae (Hemiptera: Heteroptera) from theAfrotropical and Neotropical regions, pp. 95-111 in Zootaxa 4768 (1) on pages 106-110, DOI: 10.11646/zootaxa.4768.1.6, http://zenodo.org/record/3777888, {"references":["Knyshov, A., Hoey-Chamberlain, R. & Weirauch, C. (2018) Comparative morphology of male genitalic structures in the minute litter bugs Dipsocoromorpha (Insecta: Hemiptera: Heteroptera). Journal of Morphology, 279, 1480 - 1517. https: // doi. org / 10.1002 / jmor. 20885"]}

Despite a recent surge of taxonomic work on the dipsocoromorphan family Schizopteridae, new genus-level taxa remain to be documented and described. The curation of Malaise and Yellow Pan Trap sample residues from Central and South America and Cameroon resulted in the discovery of five undescribed species that based on combined molecular and morphological phylogenetic analyses represent four phylogenetically isolated lineages within non-hypselosomatine Schizopteridae. To accommodate these new species, we here describe the four new genera Caucanannus gen. n. (Colombia, Ecuador, and Brazil) with Caucanannus perplexus sp. n. and Caucanannus novissimis sp. n.; Kamakonocoris gen. n. (Cameroon) with Kamakonocoris carinata sp. n.; Perittonannus gen. n. (Costa Rica) with Perittonannus antiquus sp. n.; and Rimanannus gen. n. (Cameroon) with Rimanannus camerunensis sp. n. The four genera are characterized by distinctive wing venation and male and female genitalic features. We provide habitus photographs and document morphological details using digital macrophotography as well as light compound microscopic and scanning electron microscopic images. Maps and measurements are also included.

Despite many bioinformatic solutions for analyzing sequencing data, few options exist for targeted sequence retrieval from whole genomic sequencing (WGS) data with the ultimate goal of generating a phylogeny. Available tools especially struggle at deep phylogenetic levels and necessitate amino-acid space searches, which may increase rates of false positive results. Many tools are also difficult to install and may lack adequate user resources. Here, we describe a program that uses freely available similarity search tools to find homologs in assembled WGS data with unparalleled freedom to modify parameters. We evaluate its performance compared to other commonly used bioinformatics tools on two divergent insect species (>200 My) for which annotated genomes exist, and on one large set each of highly conserved and more variable loci. Our software is capable of retrieving orthologs from well-curated or unannotated, low or high depth shotgun, and target capture assemblies as well or better than other software as assessed by recovering the most genes with maximal coverage and with a low rate of false positives throughout all datasets. When assessing this combination of criteria, ALiBaSeq is frequently the best evaluated tool for gathering the most comprehensive and accurate phylogenetic alignments on all types of data tested. The software (implemented in Python), tutorials, and manual are freely available at https://github.com/AlexKnyshov/alibaseq.

The genus Voragocoris Weirauch, 2012 is composed of Voragocoris schuhi Weirauch, 2012 and Voragocoris amrishi Makhan, 2013, previously recorded from Peru and Suriname, respectively. In this paper, a new species, Voragocoris weirauchae sp.n., is described based on specimens collected in the Brazilian Amazon, representing the first record of the genus from Brazil. We provide diagnosis, description, photographs of habitus, and scanning electron micrographs of the diagnostic features. A key to species based on males of Voragocoris is also presented.

Insect male genitalia show an evolutionarily variable morphology that has proven to be valuable for both, species identifications and phylogenetic analyses at higher taxonomic levels. Accurate usage of genitalic characters in taxonomic descriptions and phylogenetic analyses depends on consistency of terminology and validity of homology hypotheses. Both areas are underdeveloped in many insect groups. We here document the morphology and advance homology hypotheses of male genitalic features for the hemipteran infraorder Dipsocoromorpha, the minute litter bugs. Genitalic structures and the pregenital abdomen in Dipsocoromorpha are strikingly modified and diverse compared to other Heteroptera. In addition to variation in the shape of phallic structures (parameres and aedeagus), minute litter bug genitalia vary in the direction and degree of asymmetry and feature a plethora of processes derived from various abdominal segments with significant variation at low taxonomic levels. Here, male genitalic structures for an extensive taxonomic sample (32 genera and 71 specimens) are documented using scanning electron and confocal microscopy, and a universal terminology for genitalic structures across minute litter bugs is established that will facilitate species discovery and evolutionary research. We conclude by proposing primary homology hypotheses across the infraorder that now can be tested in a phylogenetic framework.

Evolutionary biologists have long been intrigued by exaggerated morphologies tied to sexual or natural selection. In insects, relatively few studies have investigated the evolution of such traits at the genus level and above and have used comparative phylogenetic methods to do so. We here investigate the interspecific evolution of head length in the minute litter bug genus Nannocoris Reuter based on the first phylogenetic hypothesis of the group (25 ingroup species, five gene regions, 3409 bp) and ancestral state reconstruction. Head lengths in this speciose genus range from approximately one sixth of the total body length to more than a quarter of the body length, while the head and mouthpart (rostrum) lengths are correlated. Different species therefore possess a markedly different reach of the rostrum when extended. The analyses show that head length evolution in Nannocoris is plastic, with head length elongations and reductions occurring in several clades, derived from ancestors with moderately elongated heads. Evidence is provided that exaggerated head lengths evolved through elongation of either the genal (pricei group) or the tip (arimensis group) region of the head. The biology of species in the genus Nannocoris is unknown, but given the lack of sexual dimorphism of head lengths, we speculate that head evolution in this genus may be driven by natural selection, potentially in the context of prey capture.