Synopsis of Schizopteridae (Hemiptera, Heteroptera, Dipsocoromorpha) from the United States, with description of seven new species from the US and Mexico

Abstract 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.


Introduction
The small true bug family Schizopteridae (Hemiptera: Heteroptera) in the infraorder Dipsocoromorpha contains approximately 355 described species (Emsley 1969, Hill 1984, McAtee and Malloch 1925, Wygodzinsky 1955. Although several species occur in temperate regions in Japan, Tasmania, and New Zealand, the great majority of species have been described from wet tropical and subtropical areas around the globe (Emsley 1969, Knyshov et al. 2016, Wygodzinsky 1951. The fauna of Schizopteridae in the United States could therefore be expected to be relatively small. Consistent with this prediction, only four species representing four genera were described during the first quarter of the 20 th century and these are currently the only species known to occur in the United States (Henry 1988(Henry , 2010. The four species appear to have relatively large distribution ranges in the eastern and southeastern United States including one that was originally described from Guatemala (Henry et al. 2010, Allen and Carlton 1989, Heidemann 1906, Hoffman et al. 2005, Roble and Hoffman 2000, Hoffman et al. 2007, McAtee and Malloch 1925. Heidemann (1906) described the monotypic genus Glyptocombus Heidemann, 1906 to accommodate Glyptocombus saltator Heidemann, 1906 described from specimens collected on Plummers Island in Maryland. This species was subsequently recorded from Arkansas, the District of Columbia, Michigan, Tennessee, Virginia and Washington DC (Henry 1988, Allen and Carlton 1989, Roble and Hoffman 2000. Heidemann (1906) noted the resemblance of this species to Hypselosoma Reuter and it is classified in the Hypselosomatinae (Emsley 1969). It has remained one of only a handful of New World genera in this subfamily (Uhler 1894, Emsley 1969, Carpintero and Dellapé 2006. The three remaining species of Schizopteridae recorded from the US belong to the Schizopterinae and are classified in the speciose genera Corixidea Reuter, 1891, Nannocoris Reuter, 1891, andSchizoptera Fieber, 1860. Distributions of species in the three genera range from the southern parts of South America to the US. In addition to the currently described species (9 in Corixidea, 12 in Nannocoris, and ~80 in Schizoptera: Emsley 1969, Leon andWeirauch 2016a,b), we examined specimens representing a large number of undescribed species from across the New World during taxonomic revisions that are ongoing (Weirauch Lab, unpublished data). Corixidea major McAtee & Malloch, 1925 was described from Clarksville in Tennessee and has been recorded from Florida, Virginia, Arkansas and Oklahoma (Hoffman et al. 2005, Henry et al. 2010. Nannocoris arenarius Blatchley, 1926 is known from Georgia, North Carolina, and Virginia (Hoffman et al. 2007), but was originally described from Florida (Blatchley 1926). Neither species is known to occur outside the US and both are clearly differentiated from congeneric species in Central America by head shape, wing type and venation (Nannocoris) and size, coloration and male genitalic structures (Corixidea). The situation is less clear for Schizoptera bispina McAtee & Malloch, 1925 that was originally described from Guatemala and recorded from Mexico (McAtee and Malloch 1925) and subsequently reported from Florida (Blatchley 1926). McAtee and Malloch (1925) in-dicated that the lateral spines on the male subgenital plate were shorter in the Mexican specimen compared to the holotype, but nevertheless treated them as conspecific. The species was originally classified in the subgenus Schizoptera (Lophopleurum) that contained six additional species from Central America and Trinidad (McAtee and Malloch 1925, Emsley 1969, Costas et al. 2015. Based on the lack of reciprocal monophyly and diagnostic features, Schizoptera (Lophopleurum) was recently synonymized with Schizoptera (Cantharocoris) (Leon and Weirauch 2017) that contains an additional six species from the Caribbean, Central and Northern South America including Trinidad (McAtee andMalloch 1925, Emsley 1969). Species in the subgenus Schizoptera (Cantharocoris) are recognized by the glabrous area associated with the scent gland groove extending to or beyond the midline of the metapleuron; in species of Schizoptera (Schizoptera) the glabrous area is restricted to the ventral margin of the scent gland groove.
We here provide an updated synopsis of the Schizopteridae occurring in the United States. We have examined >8,000 specimens of New World Schizopteridae as part of a project on the biodiversity and systematics of this group. The majority of specimens were curated from ethanol-preserved residues of passive trap samples, but we also borrowed point and card-mounted specimens from various collections. The 178 U.S. specimens of Schizopteridae that we have located and examined are mostly pointmounted and were borrowed from seven collections. Texas A&M and the Florida State Collection of Arthropods provided the bulk of the material, with 89 and 53 specimens, respectively. We also examined specimens of Corixidea, Nannocoris, and Schizoptera (Cantharocoris) from Mexico and other countries in Central America to assure that our synopsis of previously described and new taxa treated in this paper is as thorough as possible with respect to distribution ranges that extend beyond the administrative borders of the U.S. Additional undescribed species of the three schizopterine genera from Mexico and Central America will be treated as part of separate, genus-focused publications. In contrast, the revision of Glyptocombus presented as part of this study is comprehensive based on the available material and includes a new species known only from Mexico. We here provide a synopsis of the ten species of Schizopteridae that occur in the US, describe six of them as new, provide habitus images and genitalic illustrations of previously described and new species, and document their distribution ranges.

Materials and method
We have examined approximately 8,000 specimens of Schizopteridae from the Nearctic and Neotropical regions as part of a US National Science Foundation project on the biodiversity and systematics of the true bug infraorder Dipsocoromorpha. Among these were 178 specimens from the United States, representing the four previously described species and several undescribed species in the genera Glyptocombus, Corixidea, Nannocoris, and Schizoptera. We surveyed point-mounted specimens belonging to these four genera from Mexico and other Central American

Imaging, dissections, measurements, databasing, and distribution maps
Habitus images were taken using a Leica DFC 450 C Microsystems system (Leica, Wetzlar, Germany) with a Planapo 1.0× objective. Images of selected morphological characters were produced on the same system with a Planapo 2.0× objective. Individual images were combined using the Leica Application Suite V4.3 software or Zerene stacker V1.02 (Zerene Systems). Images were edited and assembled into image plates in Adobe Photoshop CS4 or CC2018.
To document male and female genitalia, the abdomen was separated from the body and cleared in hot 10% KOH. The male abdomen was temporarily mounted in glycerin on microscope slides, while the female abdomen was stained using Chlorazol Black E for 3 intervals of 30 seconds before slide mounting. Line drawings were prepared using a Nikon Eclipse 80i compound microscope (Nikon, Tokyo, Japan) with camera lucida.
Measurements are in mm (see Table 1; selected measurements and ratios are in text) were taken from the habitus images. Total length was measured from the anterior tip of the head to the tip of the apex of forewing, body length was measured from the anterior tip of the head to the apex of the abdomen, and width between eyes was measured in frontal perspective.
Unique specimen identifier (USI) matrix code labels with prefix and eight-digit number were associated with each specimen. Specimen information was databased using the American Museum of Natural History's Arthropod Easy Capture (AEC) database (http://www.research.amnh.org/pbi/locality/index.php). Specimen information is also available through the Heteroptera Species Pages (http://research.amnh.org/pbi/ heteropteraspeciespage/) and the http://www.discoverlife.org/ website. Maps were prepared using SimpleMappr (http://www.simplemappr.net/) from coordinates exported from the AEC database and edited using Photoshop CS4.

Nomenclatural acts
This publication and the nomenclatural acts it contains have been registered in Zoo-Bank, the online registration system for the ICZN. The ZooBank LSIDs can be resolved by appending them to the Web address http://zoobank.org/. The LSIDs for nomenclatural acts can be found in corresponding sections of this article. A larger comparative survey of wing venation across Schizopteridae (Weirauch lab, unpublished data) has indicated that the terminology used in Hypselosomatinae and Schizopterinae is inconsistent. We here introduce a slightly modified terminology for wing veins in Hypselosomatinae from those in Hill (2013) and Hoey-Chamberlain and  that also differs from the terminology used for Schizopterinae in Weirauch and Frankenberg (2015) and Leon and Weirauch (2016a,b), but is largely consistent with the terminology for Ogeriinae used by Knyshov et al. (2016). The wing vein terminology is illustrated in Figure 2 for Hypselosomatinae and in Figs 6, 8 for Schizopterinae. More extensive documentation of schizopterid forewing veins is forthcoming as part of a combined morphological and molecular phylogenetic analysis (Knyshov et al., in prep.).

Abbreviations and
In several recent publications, we have referred to the ventral sclerite of the pregenital abdomen as "sternite," following, e.g., the terminology used by Emsley (1969). Here we use the terms "tergum" and "sternum" for all dorsal and ventral abdominal sclerites and "laterotergite" for a lateral subdivision of the tergum following recent papers on the morphology of the pregenital abdomen in Dipsocoromorpha ) and a putatively closely related group of Heteroptera (Davranoglou et al. 2017).

5
Labium truncated, pronotal collar absent (Fig. 4)  Head elongated, labium very slender, posterolateral spine on metepisternum absent, R1 obliquely approaching costal forewing margin (Fig. 6)  Type species. Glyptocombus saltator Heidemann, 1906 Revised diagnosis. Recognized among genera of New World Hypselosomatinae by dense setation on all surfaces of the tibia, extending from base to apex, first labial segment not dorsally expanded, and third and fourth labial segments without processes. (Figure 1): macropterous or with elytriform forewing, length: 1.14-1.74 mm; body ovoid. Coloration: general coloration tan to black. Surface and Vestiture: scape with 5 setae, clypeus with 3 long, stout setae, buccula and first labial segment each with pair of stout ventrolateral setae, second labial segment with pair of lateral setae (Figure 1), long, stout seta dorsoapically on third tarsal segment of all legs, anophore without distinctive setae ( Figure 3A). Structure: Head: first labial segment not enlarged, clypeus simple. Thorax: tarsal formula 2-3-3, parempodia long and thin, bladder-like arolium absent. Abdomen: abdomen with 6 spiracles on left and 5 on right side, right side of tergum 8 with small projection containing spiracle, left side with large mitten-shaped projection with spiracle and long seta, sterna 7 and 8 distinct, posterior margin of sternum 7 simple, left side with triangular flap, sternum 8 rectangular-ovoid ( Figure 3B). Genitalia ( Figure 3A): right paramere long, stout with rounded apex and short, rounded process at base, left paramere scapulashaped with large thumb-shaped projection, vesica of average length with one coil and small rounded conjunctival appendage at base, anophoric process crab claw-shaped, extending posteriorly on left side.
Notes. This is the only genus of Hypselosomatinae with almost exclusive Nearctic distribution based on locality records of the single previously described and three new species, as well as one female specimen from Chiapas that represents an undescribed species. Specimens have been collected using Berlese extraction, hand collecting, yellow pan traps, pitfall traps (single and array, normal and dung), sifting leaf litter, suction traps, and UV lighting. Extracted substrates included beech humus, bottomland hardwood remnant, fallen leaves, "rubbish," earth, and the forest floor of Palmetto-gumbo limbo upland.

Diagnosis.
Recognized among species of Glyptocombus by the macropterous male, contrasting dark brown and pale legs, desclerotized portion of C+Sc vein basal to junction with R1, Cu touching M at basal corner of dc1, part of Cu distal to tc s-shaped, R2 slightly sigmoid, rc and dc approx. as long as bc (Figure 2A). Most similar to G. suteri based on wing type in males, but differentiated by details of the wing venation.
Description. Male (Figure 1): macropterous, length: 1.23 mm; body ovoid. Coloration: head, pronotum, and forewings dark brown to black, scutellum dark brown with pale lateral and apical marks, coxae, femora, and base of tibiae dark brown, apex and base of tarsi pale to yellow, pretarsi brown, genital capsule and genitalia dark brown. Surface and Vestiture: in frontal orientation forefemur without anterior stout, long seta, hind tibia with 5 erect medium-length stout setae ventrally on distal half, second tarsomere of hind leg with stout seta anteroventrally (Figure 1). Structure. Head: disc steeply decurrent anteriorly. Thorax: ratio of pronotal collar length to pronotum length 0.23, collar depressed below rest of pronotum (Figure 1), postnotum subrectangular (i.e., freely projecting portion is straight along most of posterior margin, curving only laterally, as in Rectilamina Hill; this can be observed only when specimen is in ethanol and wings have been moved aside or removed), ratio of height of forefemur to length of forefemur 0.24, ratio of length of hind tibia to width of pronotum 0.88, venation of macropterous forewing as in Figure 2A, desclerotized portion of C+Sc vein basal to junction with R1, Cu touching M at basal corner of dc1, part of Cu distal to tc s-shaped, R2 slightly sigmoid, rc and dc approx. as long as bc (Figure 2A). Abdomen and genitalia: as in genus description ( Figure 3A, B).
Female. Unknown. Etymology. Named for the collector of both known specimens of this species, Susan Halbert.
Distribution. Known only from Collier County in Florida (USA).
Description. Male (Figure 1): with elytriform forewing, length: 1.18-1.74 mm; body ovoid. Coloration: head and body dark brown to black, coxae light brown, rest of legs pale to yellow, genital capsule and genitalia light to dark brown. Surface and Vestiture: in frontal orientation forefemur without anterior stout seta, hind tibia withtwoerect medium-length setae ventrally on distal half, second tarsomere of hind leg with stout seta anteroventrally (Figure 1). Structure. Head: disc declivous. Thorax: ratio of pronotal collar length to pronotum length 0.30-0.32, pronotal collar slightly depressed below pronotum (Figure 1), posterior margin of postnotum curved, ratio of height of forefemur to length of forefemur 0.20-0.23, ratio of length of hind tibia to width of pronotum 1.00, wing venation as in Figure 2B, forewing elytriform, with veins wider than cells. Abdomen and genitalia: as in genus description ( Figure 3A, B).
Female (Figure 1): with elytriform forewing, length: 1.51 mm; body ovoid. Coloration: similar to male. Surface and vestiture: forefemur with anterior stout seta, hind tibia with four erect medium-length macrosetae ventrally on distal half, second hind tarsomere with two stout setae anteroventrally (Figure 1), entire surface of sternum 7 shiny with long setae (Figure 1). Structure. Head: disc decurrent anteriorly. Thorax: similar to male, but ratio of pronotal collar length to pronotum length 0.28, collar slightly depressed below pronotum (Figure 1), postnotum short and rectangular, ratio of height of forefemur to length of forefemur 0.26, ratio of length of hind tibia to width of pronotum 1.00, forewing similar to male. Abdomen and Genitalia: as in genus description ( Figure 3C, D).
Etymology. Named for the country of origin. Distribution. Specimens of this species have been collected in Quintana Roo and Tamaulipas in Mexico. Figures 1-3 Diagnosis. Recognized among species of Glyptocombus by reddish-brown coloration, elytriform forewings in both sexes, forewing veins narrower than cells, and areolate forewing cells (Figure 1). Most similar to G. mexicanus based on wingtype in both males and females, but differentiated by much narrower veins. (Figure 1): with elytriform forewing, length: 1.14-1.52 mm; body ovoid. Coloration: head and body reddish dark brown, legs light brown with apex of foretibiae light, genital capsule and genitalia light to dark brown. Surface and vestiture: in frontal orientation forefemur without anterior stout seta, hind tibia with four erect medium-length stout setae ventrally on distal half, second tarsomere of hind leg with stout seta anteroventrally. Structure. Head: disc decurrent anteriorly. Thorax: ratio of pronotal collar length to pronotum length 0.25-0.35, collar slightly depressed below pronotum (Figure 1), posterior margin of postnotum curved in medially around contour of scutellum, ratio of height of forefemur to length of forefemur 0.20-0.23, ratio of length of hind tibia to width of pronotum 0.94-1.16, venation of elytriform forewing as in Figure 2A, veins narrower than cells, cells areolate. Abdomen and genitalia: as in genus description ( Figure 3A, B).

Revised description. Male
Female (Figure 1): elytrous, length: 1.46-1.49 mm; body ovoid. Coloration: similar to male. Surface and vestiture: in frontal orientation forefemur without anterior stout seta, hind tibia with four erect medium-length stout setae ventrally on distal half, second hind tarsomere with stout seta anteroventrally, sternum 7 with posteriad-pointing medial U-shaped shiny area with long setae. Structure. Head: disc steeply declivous anteriorly. Thorax: similar to male, ratio of pronotal collar length Diagnosis. Recognized among Glyptocombus species by males macropterous, general coloration light brown (Figure 1, G. suteri), C+Sc not desclerotized basal to junction with R1, Cu separated from M by m-cu cross vein, Cu distal to tc straight, R2 straight, rc and dc longer than bc ( Figure 2D). Most similar to G. halbertae based on wing type in males, but differentiated by details of wing venation.
Description. Male (Figure 1): macropterous, length: 1.47 mm; body ovoid. Coloration: head and body tan, legs from middle of tibiae to pretarsus pale. Surface and Vestiture: in frontal orientation forefemur without anterior stout, long seta, hind tibia with 5 erect medium-length stout setae ventrally on distal half, second tarsomere of hind leg with stout seta anteroventrally (Figure 1). Structure. Head: disc declivous. Thorax: ratio of pronotal collar length to pronotum length 0.26, collar at same level as pronotum (Figure 1), postnotum trapezoidal, ratio of height of forefemur to length of forefemur 0.26, ratio of length of hind tibia to width of pronotum 0.94, venation of forewing as in Figure 2D, veins wider than cells. Genitalia: as in genus description ( Figure 3A, B). Notes. We have examined one undescribed female specimen of Glyptocombus from Chiapas in Mexico that has remained unassociated with male specimens (Figure 1; UCR_ENT 00094274). This specimen differs from females of Glyptocombus mexicanus sp. n. in being macropterous (elytriform forewings in males and females of Glyptocombus mexicanus sp. n.). Although wing polymorphism is common in Hypselosomatinae, species with elytriform forewings in males have not been documented to contain macropterous females (Hill 1984, Hoey-Chamberlain and Diagnosis. Recognized among species of Corixidea by the uniformly dark coloration, macropterous forewing without pale band, presence of laterally directed tergum 8 process, and very small and rounded anophoric process. Revised Description. Male (Figure 4): macropterous, body length 1.36-1.58 mm; body elongate-oval. Coloration: general coloration dark brown to black, head, thorax, and costal area of the forewing dark brown to black, remainder of wing brown to dark brown, antenna and legs uniformly brown, abdomen dark brown to black. Surface and vestiture: head, pronotum and wing veins with short to medium-length light brown setae, abdomen densely covered with elongate, light brown setae, wing cells with distinct cell-like sculpture. Structure: Head: rounded in ventral view, width of eye ~1/3 width of synthlipsis, ocellus roughly equal in size to one ommatidium, positioned close to margin of compound eye. Thorax: pronotum trapezoidal, with posterior margin slightly curved posteriorly, scutellum triangular, with rounded tip, metepisternum with posterior margin rounded, blunt metasternal process widened apically, hind coxa with well-developed adhesive pad, tarsal formula 3-3-3, pretarsus with setiform parempodia, arolia present on fore-and midleg. Abdomen: with 5-6 visible sterna corresponding to segments 2+3 (if visible), 4, 5, 6, 7, and 9 (pygophore), sternum 2 longer than preceding sterna, weakly asymmetrical, terga of pregenital abdomen moderately sclerotized, tergum 7 rectangular or weakly trapezoidal, tergum 8 roughly rectangular with process and serrated patch on left side, process of tergum 8 with narrow base and widened apex, aligned with serrated patch on tergum, bearing spinous process. Genitalia: pygophore weakly asymmetrical, dorsoventrally flattened, with rounded apex, plane of parameres rotated at 90 degrees with respect to longitudinal body axis, right paramere straight, with robust base and elongate stem, left paramere much smaller than right, roughly as long as wide, aedeagus devoid of large conjunctival appendages, vesica forming 1¼ coils, with long slender process at midpoint of vesica, anophore well sclerotized and with small rounded process.
Notes. Distinguished from other US Schizopteridae by a combination of relatively large size, blunt 3-segmented labium, absence of pronotal collar, presence of two trian- gular cells on costal margin of forewing (bordered by C+Sc, R+M, and R1, and C+Sc, R1, and R2), dorso-ventrally flattened pygophore with rounded apex, very long right paramere, and vesica with long subapical process. The genus Corixidea is a part of the Corixidea genus group, informally created by Emsley (1969), which also includes Hoplonannus McAtee & Malloch, 1925, Membracioides McAtee & Malloch, 1925, Oncerodes Uhler, 1894, Voccoroda Wygodzinsky, 1950, and Voragocoris Weirauch, 2012. Monophyly of the group is supported by the blunt labium, absence of a pronotal collar, venational similarities, and by limited molecular data when representatives of three genera were analyzed (Knyshov et al. 2016). However, generic limits that were proposed based on morphological features are unclear when undescribed diversity of the group and the poorly described original species are considered. We are currently working on a revision of the Corixidea genus group (Knyshov et al. unpublished), which will contain a comprehensive diagnosis of C. major in view of potential taxonomic changes within the group.  Reuter, 1891 and Schizoptera (Nannocoris) tuberculifera Reuter, 1891. Whereas Schizoptera is feminine, Nannocoris is masculine, because "coris" is Greek for "bug" and a noun with masculine gender. When McAtee and Malloch (1925) elevated Nannocoris to genus level, they failed to adjust the gender of previously described species from feminine to masculine and also treated their newly described species as feminine. Nevertheless, subsequent authors continued to treat Nannocoris as feminine. Nannocoris arenaria Blatchley, 1926 was corrected to Nannocoris arenarius Blatchley, 1926by Henry (1988 without comment. We here adjust the genders to masculine for all remaining names that were originally treated as feminine and are not patronyms: N. nebuliferus Reuter, 1891;N. tuberculiferus Reuter, 1891;N. capitatus (Uhler, 1894); N. nasus McAtee & Malloch, 1925;N. flavomarginatus McAtee & Malloch, 1925. Phylogeny and classification. A recent molecular phylogenetic analysis of Nannocoris (Frankenberg et al. 2018) included one North American specimen (UCR_ENT 00094252), a female from Texas that we have assigned to the new species N. brevipilus below. This taxon was recovered as part of the pricei species group that is diagnosed by the opening of the male-specific vertex gland being located in a posterior position, i.e. posteriorly on the vertex or on the pronotal collar. The opening is more anterior on the head in males of all other Nannocoris species we have examined (data not shown; see Figure 1 in Frankenberg et al. [2018] for selected species) and it is absent in some species. Males of N. arenarius and the second newly described species, N. anophorus, share the pronotal position of the opening of the vertex gland. We therefore tentatively assign all three Nearctic species to the pricei species group. Diagnosis. Recognized among species of Nannocoris by relatively short head, ovoid body and forewing shape, yellow costal and posterior claval margins, long setae on forewing veins, vertex gland opening on depression of pronotal collar, short, smoothly rounded vesica with less than one coil, apically bifurcating right paramere, and long and sigmoid anophoric process that reaches anteriorly to terga 5 or 6.

Female: Unknown.
Etymology. Named for the long anophoric process that is unusually prominent among species of Nannocoris, especially in other species of the pricei species group.
Notes. The holotype was collected using a flight intercept trap. Distribution. Known only from Hays County in Texas. Figures 6, 7, 10 Material. Type material. Holotype: male: USA: Florida: Pinellas Co.: Dunedin, 28.027°N 82.77126°W, Jan 4-Feb 16, W.S. Blatchley (PURC). Paratypes: same data as holotype, 11 specimens. Revised diagnosis. Recognized among species of Nannocoris by relatively short head, ovoid body and forewing shape, yellow costal and posterior claval margins, short setae on forewing veins, vertex gland opening on depression of pronotal collar, short and thin, slightly sigmoid vesica, apically bifurcating right paramere, and relatively short s-shaped anophoric process with slender base that reaches anteriorly to tergum 7. Similar to N. brevipilus, but distinguished by slender base of anophoric process.
Diagnosis. Recognized among species of Nannocoris by relatively short head, ovoid body and forewing shape, yellow costal and posterior claval margins, short setae on forewing veins, vertex gland opening on depression of pronotal collar, short and thin, slightly sigmoid vesica, apically bifurcating right paramere, and relatively short s-shaped anophoric process with slender base that reaches anteriorly to tergum 7. Similar to N. brevipilus, but distinguished by slender base of anophoric process.
Female ( Figure 6): similar to male, length: 1.11-1.23 mm, shorter and more ovoid than male, yellow borders of costal and claval margin less pronounced, forewings shorter.
Etymology. Named for the short vestiture on the wings that distinguishes this species from the second species occurring in Texas, N. anophorus; a combination of the Latin brevis (short) and pilus (hair).
Notes. We treat a series of female specimens collected in Bastrop County as conspecific with N. brevipilus. We refrain from treating these specimens as paratypes, because no syntopic males are available and females of N. anophorus are unknown. We argue that these specimens are unlikely to be conspecific with N. anophorus because of the short vestiture, but we cannot exclude the possibility that they are females of a yet undiscovered species of Nannocoris.
Males were collected using pitfall and flight intercept traps, females with Berlese extraction.
Female. One female specimen reported (but not illustrated) by Blatchley (1926) and not examined in our study.
Notes. McAtee and Malloch (1925) described this species from Guatemala based on a single male specimen. They mentioned the slightly yellowish humeral angle, dark color proximally across the membrane, and only slightly notched scutellar apex as characters distinguishing this species from Schizoptera (Cantharocoris) sulcata McAtee & Malloch, 1925, while emphasizing the importance of the shape of the subgenital plate. The characteristic lateral spines on the subgenital plate are shorter in the specimen from Mexico that McAtee and Malloch (1925) considered conspecific with S. (C.) bispina. Blatchley (1926) reported S. (C.) bispina from Florida and provided a redescription. The specimens examined by us are clearly conspecific with those examined and illustrated by Blatchley (1926) based on his fairly detailed description of coloration and drawing of the subgenital plate. No other described species of Schizoptera (Cantharocoris) has a subgenital plate that even remotely resembles the one in S. (C.) bispina. Nevertheless, the distribution range of S. (C.) bispina is much larger than the ranges typically seen in schizopterids. A comprehensive revision of Schizoptera (Cantharocoris) across the Nearctic and Neotropical regions is therefore not unlikely to reveal that the current concept of S. (C.) bispina is a complex of several closely related species. Blatchley (1926) reported the three specimens he examined as "beaten from Spanish moss," "sifted from vegetable debris," and "beaten from sugar cane." New records indicate that specimens were collected by UV lighting and Malaise trapping.
Distribution. Guatemala, Mexico, and Florida, Louisiana, and Texas in the United States.

Description. Male
Female (Figure 8): coleopteroid with very narrow membranous border, length: 1.22-1.29 mm; body ovate. Coloration: (Figure 8): more uniformly brown compared to male, head slightly paler, legs pale yellow, An2 anteriorly and claval furrow lined with contrasting dark coloration, similar to pattern in male. Surface and vestiture: similar to male. Structure: Head: triangular in frontal view, approx. as wide as high (Figure 8), synthlipsis slightly more than 3 times width of eye. Thorax: pronotum narrower than in male, posterior pronotal margin medially slightly concave, forewing veins obsolete, claval furrow distinct, posterior margin of clavus broadly elevated. Genitalia: not examined.
Etymology. Named in honor of Dr. Ed Riley, who collected most of the specimens examined for this study. A noun in genitive case.
Notes. Coleopteroid females evolved several times independently in the Schizoptera genus group (Leon and Weirauch 2017), but female coleopteroidy appears to be especially prevalent in the subgenus Schizoptera (Cantharocoris). Associating conspecific macropterous males and coleopteroid females is often a challenge in Schizoptera. Species diagnoses heavily rely on male genitalic features, long series of males and females from a small collection event are rare, and molecular matching of males and females has been done only for a few minute litter bugs (Knyshov et al. 2016). Matching male and female Schizoptera rileyi was comparatively straightforward because of the long series of specimens of both sexes from across the known distribution range, but also because of the distinctive dark lines on the forewings.
Although the collection method is unknown for the majority of specimens examined, both sexes have been collected using flight intercept traps and some males have been taken at UV light traps.
Distribution. Known only from the U.S., where it has been collected in Natchitoches Parish in Louisiana and Brazos, Cameron, and Hidalgo Counties in Texas.
Etymology. Named in honor of Dr. Thomas Henry on the occasion of his 70 th birthday and in recognition of his outstanding contributions to heteropterology. We also thank Tom for advancing our understanding of the little-studied Schizopteridae in the United States by keeping his dedicated eye open for these tiny bugs and publishing updated distribution records. A noun in genitive case.
Distribution. Known from Tamaulipas and Veracruz in Mexico and Cameron County in Texas in the U.S.
Notes. This species is unique among Schizoptera in the U.S. in having brown and yellow contrasting legs, in addition to the distinctive features of the male abdomen and genitalia.