REVISION AND PHYLOGENY OF THE FERN-INHABITING GENUS FELISACUS DISTANT (INSECTA: HETEROPTERA: MIRIDAE: BRYOCORINAE)
ANNA A. NAMYATOVA AND GERASIMOS CASSIS
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY REVISION AND PHYLOGENY OF THE FERN-INHABITING GENUS FELISACUS DISTANT (INSECTA: HETEROPTERA: MIRIDAE: BRYOCORINAE)
ANNA A. NAMYATOVA AND GERASIMOS CASSIS Evolution and Ecology Research Centre, School of Biological, Earth and Environmental Sciences, University of New South Wales
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 403, 168 pp., 22 figures, 1 table Issued June 22, 2016
Copyright © American Museum of Natural History 2016 ISSN 0003-0090 CONTENTS Abstract ...... 4 Introduction ...... 4 Material and methods ...... 5 Specimens ...... 5 Terminology ...... 5 Microscopy and imaging ...... 5 Phylogenetic analysis ...... 6 Morphology ...... 6 Results ...... 7 Phylogeny ...... 7 Discussion of major nodes ...... 18 Taxonomy ...... 25 Felisacus ...... 25 Checklist of Felisacus species ...... 28 Key to Felisacus species ...... 29 Felisacus adamsi ...... 34 Felisacus albus ...... 35 Felisacus amboinae ...... 40 Felisacus auritulus ...... 51 Felisacus australicus ...... 52 Felisacus bau ...... 55 Felisacus bellus ...... 57 Felisacus bismarckensis ...... 59 Felisacus bradi ...... 61 Felisacus caledonicus ...... 63 Felisacus ceylonicus ...... 65 Felisacus crassicornis ...... 67 Felisacus cristobalus ...... 68 Felisacus curvatus ...... 70 Felisacus dauloi ...... 74 Felisacus elegantulus ...... 77 Felisacus fedori ...... 80 Felisacus filicicola ...... 82 Felisacus glabratus ...... 85 Felisacus gressitti ...... 87 Felisacus indicus ...... 89 Felisacus insularis ...... 90 Felisacus jacobsoni ...... 91 Felisacus javanus ...... 95 Felisacus lambkinae ...... 98 Felisacus linae ...... 100 Felisacus lindbergae ...... 103 Felisacus liui ...... 105 Felisacus longiceps ...... 107
2 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 3
Felisacus lordhowensis ...... 110 Felisacus luzonus ...... 112 Felisacus madagascariensis ...... 116 Felisacus magnificus ...... 117 Felisacus malayensis ...... 121 Felisacus meilingae ...... 122 Felisacus minutus ...... 125 Felisacus myersi ...... 126 Felisacus nigrescens ...... 129 Felisacus nigricornis ...... 131 Felisacus ochraceus ...... 132 Felisacus ovalau ...... 134 Felisacus philippinensis ...... 135 Felisacus ponaponensis ...... 138 Felisacus rubricuneus ...... 139 Felisacus schuhi ...... 141 Felisacus senaru ...... 143 Felisacus signis ...... 145 Felisacus solomonicus ...... 146 Felisacus tanna ...... 148 Felisacus usingeri ...... 150 Felisacus vitilevu ...... 151 Felisacus wangae ...... 154 Felisacus webbi ...... 156 Felisacus yasunagai ...... 157 Felisacus zuparkoi ...... 159 Acknowledgments ...... 161 References ...... 162 Appendix 1: Characters and character states used in the phylogenetic analysis ...... 165 Data matrix ...... 168 ABSTRACT The fern-feeding genus Felisacus (Miridae: Bryocoronae) was revised. The genus contains 55 spe- cies, 29 are new to science, they are: Felisacus albus, F. australicus, F. bau, F. bismarckiensis, F. bradi, F. caledonicus, F. ceylonicus, F. cristobalus, F. fedori, F. indicus, F. lamkinae, F. linae, F. lindbergae, F. liui, F. lordhowensis, F. luzonus, F. malayensis, F. meilingae, F. myersi, F. ovalau, F. schuhi, F. senaru, F. solomonicus, F. tanna, F. vitilevu, F. wangae, F. webbi, F. yasunagai, F. zuparkoi. Felisacus capitatus is synonymized with F. magnificus; F. okinawanus is synonimized with F. longiceps F. carpenterae is synonymized with F. javanus. Felisacus jacobsoni, previously synonymized with F. javanus, is treated as a separated species. The status of F. signis, described as a subspecies of F. carpenterae, is raised to a separate species. Genus Felisacoris is synonymized with Felisacus. The diagnosis and key are pro- vided for all species and descriptions are provided for the species, where the specimens were avail- able. The drawings of male genitalia are provided where available, and drawings of female genitalia are given for some species. Phylogenetic analysis with inclusion of most of the species has been performed and discussed. Keywords: systematics, taxonomy, species, description, diagnosis, key, fern-feeding
INTRODUCTION Motschulsky (1863) described Liocoris Motschulsky, 1863, for a single species, L. glabratus Felisacus Distant, 1904, belongs to the (Motschulsky, 1863), in the suprageneric division hyperdiverse family Miridae (Insecta: Heter- Capsides (= Miridae). Liocoris was preoccupied optera), and is comprised of more than 11,000 and Distant (1904) gave Felisacus as a replacement species (Cassis and Schuh, 2012). The genus belongs to the morphologically heterogeneous name and transferred it to the family Capsidae, subfamily Bryocorinae, for which we recently subfamily Mirinae, division Cylaparia. Reuter erected a new tribe—the Felisacini (Namya- (1904) described Hyaloscytus Reuter, 1904, which tova et al., 2016; Namyatova and Cassis, 2016). was synonymized with Felisacus by Poppius (1911). Felisacus has an Indo-Pacific distribution Prior to this work the genus comprised 26 species. (Schuh and Stonedahl, 1986), a pattern that is Woodward (1954, 1958) treated a majority of Feli- repeated for other mirid genera, including sacus species and recognized two informal species Helopeltis Signoret, 1858 (Stonedahl, 1991), groups: “magnificus group” and “elegantulus Coridromius Signoret, 1862 (Tatarnic and Cas- group,” based on external morphology and the sis, 2010), Ragwelellus Odhiambo, 1962 shape of the parameres. (Namyatova and Cassis, 2016), Peritropis Other works of significance, inlcude Carv- Uhler, 1891 (Gorcyza, 2000; Moulds and Cas- alho’s (1956, 1981) treatment of the Microne- sis, 2006) and Pseudoloxops Kirkaldy, 1905 sian and Papua New Guinean species; Hu and (Schuh, 1995; Schuh, 2002–2013). Zheng (2001), Lin (2000), Miyamoto (1965), Felisacus is noteworthy because of its asso- and Usinger (1946) described new species ciations with ferns, which is atypical for insects from China, Taiwan, Japan, and Guam respec- (e.g., Weintraub et al., 1995; Jensen and Hol- tively; and Hsiao (1944) described new species man, 2000; Konstantinov and Knyshov, 2015; based on material housed in the United States Schneider, 2016). Within the Miridae, fern- National Museum. associated taxa include Felisacus, species of the The aim of this paper is to provide a detailed tribe Bryocorini (Konstantinov and Knyshov, diagnosis and redescription of the genus and its 2015) and a single Macrolophus Fieber, 1858, included species, redescription of numerous species which belongs to the tribe Dicyphini new species, as well as determining the phylo- (Wheeler et al., 1979). genetic relationships of the species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 5
METHODS AND MATERIALS UNSW University of New South Wales (Sydney, Australia) Specimens UCB University of California, Berkeley Around 800 specimens were examined for the USNM United Stated National Museum, Smith- present study and were borrowed from the fol- sonian Institution (Washington, D.C.) lowing collections: WAMP Western Australian Museum (Perth, AM Australian Museum (Sydney, Australia) Australia) AMNH American Museum of Natural History ZISP Zoological Institute, Russian Academy (New York) of Sciences (St. Petersburg, Russia) ANIC Australian National Insect Collection A unique specimen identifier (USI) with a (Canberra, Australia) matrix code was attached to each specimen. BMNH Natural History Museum (London, Those codes have a AMNH_PBI prefix, unless UK) otherwise stated. All collection-event data and BPBM Bernice P. Bishop Museum (Honolulu) host-plant information were entered in the Plant CAS California Academy of Sciences (San Bug Planetary Biodiversity Inventory Locality Francisco) Database (https://research.amnh.org/pbi/local- HNHM Hungarian Natural History Museum ity/), and these data are also served on the Dis- IOZ Institute of Zoology, Chinese Academy cover Life website (http://www.discoverlife.org/) of Sciences (Beijing, China) and Heteroptera Species Pages (http://research. amnh.org/pbi/heteropteraspeciespage/). ISNB Royal Institute of Natural Science of Maps: Coordinates were mapped using Arc- Belgium (Brussels, Belgium) GIS (https://www.arcgis.com/features/), using MNHN National Museum of Natural History the ESRI World Physical Map as a background. (Paris, France) Terminology: Terminology of male genitalia MVMA Museum of Victoria (Melbourne, follows Kerzhner and Konstantinov (1999) (see Australia) fig. 10B, C for structures’ names) with aedeagus MZH Finish Museum of Natural History divided into conjunctiva and vesica (= endosoma (Helsinki, Finland) of Cassis, 2008). The terminology of female geni- NHRS Naturhistorika riksmuseet (Stockholm, talia follows Davis (1955) and Schwartz (2011). Sweden) Microscopy and imaging: Genitalia were dis- NKMU Nankai University Insect Collection sected using Leica MZ 16 microscope in the Uni- (Tianjin, China) versity of New South Wales and Nikon SMZ 1500 NML Nationaal Natuurhistorische Museum microscope in the Saint Petersburg State University. (Leiden, the Netherlands) Most dorsal habitus images were taken at the Uni- NMNS National Museum of Natural Science versity of New South Wales using a Visionary Digi- (Taichung, Taiwan) tal BK Plus Lab photographic system (www. NTM Northern Territory Museum (Darwin, visionarydigital.com) with a Canon EOS 40D cam- Australia) era. Images of type specimens of Felisacus auritulus QM Queensland Museum (Brisbane, Distant, 1913, were taken in Sackler’s Biodiversity Australia) Imaging Lab at the Natural History Museum, Lon- SAMA South Australian Museum (Adelaide, don, using a Canon 450D camera attached to a Australia) Leica MZ 9.5 stereomicroscope. Images of the spec- TAMU Texas A&M University (College Stat- imens of F. adamsi Carvalho, 1956, Felisacus fedori, ion, Texas) sp. nov., F. senaru, sp. nov., F. carpenterae Hsiao, TYCN Tomohide Yasunaga personal collection 1944, F. ponaponensis Carvalho, 1956, F. rubricu- (Nagasaki, Japan) neus Carvalho, 1956, F. signis Hsiao, 1944, and F. 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 webbi, sp. nov., and male of F. jacobsoni Poppius, Forty-eight species of Felisacus were ana- 1914, were made in the Department of Entomol- lyzed. We did not include F. nigricornis Poppius, ogy, Saint Petersburg State University using Nikon 1912, as specimens were unavailable for study. D700 SLR digital camera attached to Nikon SMZ We also excluded F. adamsi, F. auritulus Distant, 1500 stereomicroscope. Multiple images of all spec- 1913, F. crassicornis Usinger, 1946, F. gressitti imens were merged using Helicon Focus software Miyamoto, 1965, F. madagascariensis Poppius, (http://www.heliconsoft.com). All images were pro- 1912, and F. usingeri Woodward, 1954, because cessed using Adobe Photoshop CS3 extended and males were unavailable. CS5.1 extended software. Four outgroup taxa included two Bryocorinae Measurements: Measurements are shown in species: Pachypeltis reuteri (Stål, 1871) and table 1 and include the following: body length, Nesidiocoris tenuis (Reuter, 1895). The trees were clypeus to cuneus length, length of antennal seg- rooted with Stenotus binotatus (Fabricius, 1794) ments I and II, head width, vertex width, prono- (Mirinae). tum length and width. Measurements were mostly taken using a digital micrometer and Morphology recorded with Winwedge software (www.win- wedge.com). Species of Felisacus all have very similar mor- All scalebars are 0.1 mm in length, unless oth- phology. There are only a few external characters erwise stated. that have diagnostic and phylogenetic value, Phylogenetic analysis: The character including: shape of head from dorsal view; shape matrix was created in Mesquite software (Mad- and relative length of antennal segment I; length dison and Maddison, 2010), and includes 72 of labium, length and shape of labial segments characters of external and genitalic morphology I–III; shape of cuneus; and coloration of anten- (see SI 1 for matrix). Characters and character nal segments I–II, pronotum, hemelytron, and states are listed in appendix 1. More than half of femora (see generic description for the details). the characters coded pertain to the male genita- The female genitalia are diagnostic for major lia (appendix 1). groups of species, but are more or less uniform All analyses were run using TNT (http://www. within these groups. In one group, the dorsal cladistics.com/aboutTNT.html) (Goloboff et al., labiate plate is very small and transparent, which 2000, 2008) and NONA (http://www.cladistics. rendered it of limited diagnostic and phyloge- com/aboutNona.htm) (Goloboff, 1993) imple- netic value. mented through Winclada (http://www.cladis- The key characters within the genus are those of tics.com/wincDownload.htm) (Nixon, 1999) the male genitalia, particularly the aedeagus and software, with parsimony as the optimally crite- parameres. These characters are variable at the sub- rion. For the analyses in TNT, we employed tra- clade level and between closely related species. The ditional searches with 10000 replications and 10 genital capsule is less variable and simple. trees saved per replications. All characters were The right parameres were drawn mostly in the treated as unweighted and unordered. Searches ventral view to demonstrate setal pattern, which were also undertaken using implied weights with is not visible in the dorsal view. We used descrip- K = 3–20. All unsupported nodes were collapsed tive terms for the right paramere, with the posi- after each analysis. tion of components referred to as basal, medial Bremer support values (Bremer, 1994) were or apical, and outer and inner angles. All of these calculated using the Bremer script in TNT. Boot- are shown on figure 11. The left paramere is illus- strap resampling (Felsenstein, 1985) based on trated in dorsal view. unweighted trees was performed as implemented Aedeagal characters were used extensively in Winclada with 10,000 replications. for species differentiation, and these included: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 7 shape of sclerite around secondary gonopore, sky, 1863), F. indicus, sp. nov., and F. insularis sclerotization of conjunctiva and number and Miyamoto, 1965 (fig. 8A, J, K, L). shape of vesical spicules. The vesica of species The spicules placed on the left-hand side have within clades 2 and 14 possesses spicules of dif- a tendency to form a concavity apically. Their ferent shape. Within clade 3, only F. zuparkoi, homologization is hampered, as those spicules sp. nov., has a small claw-shaped spicule. Within vary in size and position, and in some cases there clade 18, comprising 30 species, the spicules are are two or more spicules with similar concavities usually distinct. within the same species (e.g., figs. 8K, M, 9D, K). In some cases we homologized the spicules However, we were able designate a spicule H in based on their position. Spicule A occurs in 21 F. australicus and F. ochraceus (figs. 8C, 9F). This species and is placed dorsally on the right-hand spicule is placed on the left-hand side adjacent to side; it is sinuate, curved dorsoventrally, and is spicule C, and has a distinct hook-shaped con- usually broad basally and elongate. In F. filicicola cavity apically, and is narrow and of moderate and F. tanna, sp. nov., spicule A has the same length. In addition, we delimit a spicule G, which position and shape, but is short (figs. 8I, 9M), is broad and long, has a distinct concavity api- whereas in F. meilingae, sp. nov., and F. myersi, cally, and is placed ventrally on the left-hand side sp. nov., it is straight (figs. 9B, C). (figs. 8E, G, 9B, C). Spicule B occurs in 21 species and is placed ventrally on the right-hand side or medially, or RESULTS dorsal with respect to other spicules; it is more or less C-shaped and usually broad medially. Phylogeny This spicule exhibits continuous variation in size The unweighted analysis in TNT produced in this group; from very short in F. amboinae (fig. 8750 equally parsimonious trees with 172 steps, 8B) to very long in F. australicus, sp. nov., and F. CI = 0.47 and RI = 0.83. The implied weight ochraceus Usinger, 1946 (figs. 8C, 9F). InF. filici- analyses with K from 3 through 20 resulted in 70 cola (Kirkaldy, 1908), F. jacobsoni Poppius, 1914, (K = 9, 19) to 811 (K = 3) trees. In the current and F. signis Hsiao, 1944, spicule B is not broad analysis, the consensus trees obtained from anal- medially (figs. 8M, 9K). yses with K = 3–6 are identical, and consensus Spicule C is present in 12 species and is placed trees with K = 7–11 are also identical and have on the left dorsal position, and is sinuate or better resolution. The analyses run with K = broad medially, and is usually dorsoventrally 16–20 were also stable, slightly differing in topol- curved basally. ogy from the trees obtained with K = 7–11 at Spicule D is small, basal, and dorsal in ori- node 18. We have chosen one of the most parsi- entation. Spicule I is elongate, broad basally monious trees obtained with the K = 7 for opti- and placed on the right-hand side, and more mization of characters. There are alternative proximal to the middle. These spicules are views on K value selected for optimization, with found only in F. meilingae and F. myersi (fig. some authors finding more-resolved trees of 9B, C). greater trees with K >5 results (Reinert et al., Spicule E is placed dorsally on the right-hand 2009; Harbach et al., 2012; Penz et al., 2013; Val- side, sclerotized apically and membranous dez-Mondragon, 2013, 2014; Namyatova et al., mediobasally. It occurs in F. filicicola, F. lordhow- 2016). Trees with K >10 are very rarely chosen ensis, sp. nov., and F. tanna (figs. 8I, R, 9M). (e.g., Namyatova et al., 2016). Our results corre- Spicule F is long, needle shaped, placed on the spond with the “plateaus of stability” found in left-hand side, is ventral in position and can be Reinert et al. (2009) and Harbach et al. (2012). obscured by other spicules in dorsal view. It Bremer supports and bootstrap support val- occurs in F. albus, sp. nov., F. glabratus (Motschul- ues are given in figure 1. 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
TABLE 1 Measurements of Felisacus spp.
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F. adamsi ♂ (N=2) F1 2.90 2.38 0.63 0.90 0.85 0.48 0.75 0.25 F2 3.10 2.33 0.65 0.90 0.78 0.48 0.78 0.25 F. albus ♂ (N=2) M1 3.72 2.55 0.71 1.02 0.83 0.58 0.87 0.24 M2 4.25 2.87 0.75 1.15 0.88 0.61 1.02 0.26 ♀ (N=1) F1 3.81 2.70 0.74 1.02 0.73 0.59 1.03 0.28 F. amboinae ♂ (N=2) M1 3.00 2.63 0.63 0.96 0.80 0.48 0.74 0.22 M2 3.34 2.61 0.66 1.09 0.90 0.55 0.76 0.24 ♀ (N=4) F1 3.43 2.91 0.70 1.07 0.83 0.52 0.76 0.24 F2 3.30 2.76 0.67 - - 0.50 0.78 0.22 F3 3.26 2.87 0.67 1.04 0.85 0.48 0.78 0.22 F4 3.26 2.83 0.70 0.96 0.85 0.50 0.76 0.24 F. auritulus Sex unknown (N=5) SU1 3.96 - 0.8 0.88 0.56 0.44 1.0 0.28 F. australicus ♂ (N=2) M1 3.91 2.79 0.76 1.19 1.00 0.61 0.92 0.27 M2 4.20 2.94 0.77 1.21 0.95 0.63 1.01 0.27 ♀ (N=1) F1 4.42 3.14 0.84 1.49 0.97 0.68 1.02 0.27 F. bau ♂ (N=1) M1 3.47 2.71 0.67 - 0.93 0.54 0.83 0.25 ♀ (N=1) F1 3.75 2.77 0.66 1.11 0.93 0.52 0.81 0.25 F. bellus ♂ (N=2) M1 4.45 3.12 0.81 1.33 0.95 0.63 1.02 0.28 M2 3.74 2.68 0.74 1.12 0.88 0.53 0.86 0.27 ♀ (N=4) F1 4.08 3.03 0.80 1.10 0.89 0.58 0.96 0.29 F2 4.04 3.01 0.82 1.11 0.80 0.61 1.04 0.30 F3 4.30 3.08 0.78 - - 0.61 1.05 0.30 F4 4.42 3.15 0.84 1.25 1.86 1.05 1.56 0.62 F. bismarckensis ♂ (N=1) M1 3.07 2.09 0.61 0.84 0.48 0.49 0.78 0.25 ♀ (N=3) F1 3.53 2.46 0.69 - 0.55 0.53 0.86 0.27 F2 3.72 2.62 0.74 0.93 0.62 0.55 0.92 0.29 F3 3.57 2.39 0.68 0.91 0.54 0.53 0.88 0.27 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 9
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F. bradi ♂ (N=1) M1 4.27 3.24 0.79 1.25 0.83 0.64 1.01 0.30 ♀ (N=1) F1 4.94 3.83 0.88 1.22 0.75 0.74 1.19 0.37 F. caledonicus ♂ (N=5) M1 3.67 2.49 0.66 1.20 1.00 0.52 0.85 0.26 M2 3.54 2.58 0.71 1.13 0.99 0.55 0.79 0.26 M3 3.71 2.56 0.69 1.14 0.98 0.54 0.81 0.25 M4 3.56 2.58 0.68 1.13 0.79 0.53 0.78 0.27 M5 3.80 2.61 0.66 1.20 0.97 0.55 0.85 0.27 ♀ (N=3) F1 3.99 2.72 0.71 1.14 0.96 0.56 0.95 0.28 F2 3.76 2.52 0.62 1.26 0.93 0.54 0.90 0.29 F3 3.76 2.59 0.70 1.20 0.97 0.56 0.88 0.28 F4 3.88 2.71 0.69 1.20 0.96 0.55 0.89 0.28 F5 3.74 2.70 0.73 1.15 0.91 0.51 0.89 0.28 F. ceylonicus ♂ (N=2) M1 2.61 2.28 0.61 0.63 0.43 0.35 0.67 0.15 M2 2.91 2.48 0.67 0.61 0.52 0.37 0.78 0.20 F. crassicornis ♀ (N=5) F1 3.35 2.33 0.68 0.75 0.57 0.60 0.89 0.33 F2 3.44 2.44 0.69 0.78 0.56 0.59 0.90 0.33 F3 3.47 2.41 0.70 0.89 0.64 0.58 0.88 0.32 F4 3.36 2.32 0.66 0.80 0.55 0.58 0.91 0.33 F5 3.31 2.50 0.68 0.86 0.66 0.59 0.93 0.31 F. cristobalus ♂ (N=1) M1 3.48 2.50 0.78 1.06 0.68 0.58 0.85 0.26 ♀ (N=2) F1 4.16 2.94 0.83 1.14 0.72 0.63 0.96 0.29 F2 4.21 2.95 0.84 1.10 0.80 0.64 0.99 0.31 F. curvatus ♂ (N=5) M1 4.19 2.86 0.91 1.04 0.75 0.54 0.95 0.25 M2 4.54 3.18 0.91 - 0.78 0.54 0.99 0.27 M3 4.45 3.20 0.98 1.08 0.79 0.56 1.05 0.28 M4 4.60 3.12 0.91 0.93 0.86 0.56 1.10 0.29 M5 4.76 3.39 1.03 1.22 0.87 0.61 1.13 0.29 ♀ (N=5) F1 5.42 3.81 1.11 1.32 0.86 0.61 1.24 0.30 F2 5.39 3.45 0.95 1.18 0.82 0.58 1.23 0.28 F3 4.37 2.98 0.80 0.99 0.80 0.54 1.06 0.27 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F4 5.17 3.53 1.09 1.22 0.85 0.61 1.23 0.31 F5 4.94 3.49 1.03 1.10 0.76 0.59 1.19 0.29 F. dauloi ♂ (N=4) M1 3.89 2.69 0.72 - - 0.65 1.04 0.30 M2 3.94 2.72 0.74 1.23 0.96 0.58 1.03 0.25 M3 3.80 2.68 0.63 - 0.92 0.61 0.99 0.28 M4 3.98 2.76 0.72 - - 0.57 0.93 0.25 ♀ (N=5) F1 4.52 3.08 0.80 1.22 0.99 0.67 1.08 0.32 F2 4.36 2.93 0.77 1.20 0.98 0.65 1.04 0.30 F3 4.51 3.02 0.80 1.25 0.93 0.58 1.03 0.25 F4 4.32 2.92 0.79 1.26 1.03 0.61 0.99 0.28 F5 - 3.04 0.85 1.35 0.19 0.57 0.93 0.25 F. elegantulus ♂ (N=5) M1 3.95 2.68 0.67 1.09 0.82 0.53 0.90 0.25 M2 4.09 2.81 0.74 1.11 0.96 0.57 0.97 0.26 M3 4.19 2.84 0.72 1.16 0.93 0.56 0.98 0.26 M4 4.18 2.81 0.73 1.13 0.96 0.53 0.99 0.26 M5 4.02 2.76 0.74 1.08 0.91 0.58 0.96 0.27 ♀ (N=5) F1 4.68 3.25 0.83 1.20 0.93 0.63 1.13 0.31 F2 4.42 2.96 0.78 1.22 0.97 0.59 1.03 0.30 F3 4.44 3.01 0.79 1.17 0.89 0.61 1.04 0.31 F4 4.08 2.78 0.71 1.13 0.92 0.59 0.98 0.29 F5 4.18 2.78 0.75 1.10 0.83 0.58 1.00 0.28 F. fedori ♂ (N=2) M1 4.46 0.81 0.92 1.42 0.81 0.71 1.10 0.32 M2 4.39 0.78 0.92 1.49 0.78 0.67 1.10 0.28 ♀ (N=3) F1 5.81 1.03 1.03 1.70 1.03 0.78 1.45 0.28 F2 5.10 0.92 0.96 1.49 0.92 0.78 1.27 0.32 F3 4.88 0.88 1.06 1.45 0.88 0.74 1.27 0.32 F. filicicola ♂ (N=5) M1 3.76 2.61 0.67 1.10 0.89 0.55 0.85 0.27 M2 3.51 2.47 0.67 1.09 0.91 0.57 0.80 0.27 M3 3.61 2.61 0.70 1.12 0.97 0.59 0.85 0.26 M4 3.73 2.63 0.69 1.12 1.00 0.55 0.86 0.26 M5 3.64 2.58 0.69 1.17 0.96 0.57 0.86 0.28 ♀ (N=5) F1 3.86 2.65 0.70 1.08 0.89 0.58 0.89 0.26 F2 3.47 2.38 0.62 0.95 0.85 0.53 0.79 0.25 F3 3.91 2.69 0.74 1.10 1.02 0.62 0.94 0.30 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 11
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F4 3.82 2.65 0.69 1.13 0.85 0.62 0.91 0.30 F5 3.46 2.51 0.72 1.09 0.89 0.62 0.85 0.30 F. glabratus ♂ (N=1) M1 3.85 2.73 0.75 - - 0.58 0.96 0.29 ♀ (N=1) F1 4.03 2.91 0.82 1.06 0.81 0.60 1.04 0.30 F. gressitti ♀ (N=1) F1 3.09 2.74 0.59 0.78 0.61 0.48 0.76 0.22 F. indicus ♂ (N=1) M1 3.96 2.74 0.75 1.05 0.81 0.61 0.93 0.24 F. insularis ♂ (N=4) M1 4.21 2.91 0.73 0.95 0.86 0.62 1.04 0.31 M2 3.73 2.59 0.68 1.10 0.90 0.50 0.81 0.24 M3 4.30 3.02 0.84 1.09 0.80 0.68 1.01 0.31 M4 3.90 2.71 0.74 1.01 0.78 0.59 0.97 0.28 ♀ (N=4) F1 4.07 2.80 0.79 1.09 0.82 0.58 0.97 0.26 F2 4.30 3.10 0.83 1.01 0.74 0.58 1.04 0.27 F3 3.82 2.69 0.76 0.98 0.77 0.55 0.83 0.26 F4 3.83 2.65 0.66 1.04 0.74 0.58 0.95 0.25 F. jacobsoni ♂ (N=2) M1 4.66 3.21 0.85 1.37 1.06 0.65 1.01 0.27 M2 4.40 3.00 0.79 1.44 1.03 0.63 1.01 0.29 ♀ (N=2) F1 4.77 3.36 0.88 1.39 0.96 0.65 1.10 0.31 F2 4.61 3.31 0.82 1.29 1.01 0.58 0.91 0.26 F. javanus ♂ (N=5) M1 3.04 2.40 0.65 - 0.82 0.51 0.71 0.20 M2 - 2.37 0.65 - 0.87 0.53 0.73 0.25 M3 3.69 2.68 0.72 1.10 0.88 0.55 0.83 0.27 M4 - 2.72 0.72 1.06 0.83 0.58 0.87 0.25 M5 3.40 2.45 0.73 - - 0.55 0.83 0.18 ♀ (N=5) F1 3.57 - 0.64 1.10 - 0.57 0.86 0.27 F2 3.54 - 0.75 - 0.87 0.59 0.89 0.25 F3 3.89 2.87 0.74 1.07 0.84 0.64 0.95 0.32 F4 4.02 2.92 0.75 - 0.87 0.58 0.91 0.27 F5 4.06 2.98 0.77 - - 0.62 0.92 0.28 F. lambkinae ♂ (N=5) M1 3.17 2.29 0.56 0.94 0.76 0.49 0.71 0.19 M2 2.50 0.56 0.96 0.62 0.51 0.72 0.21 . M3 3.14 2.36 0.62 0.90 0.73 0.47 0.69 0.18 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F1 3.45 2.41 0.59 0.94 0.81 0.53 0.79 0.26 F2 3.18 2.37 0.63 0.95 0.73 0.52 0.76 0.24 F3 2.57 0.65 0.92 0.77 0.49 0.83 0.24 F4 3.33 2.42 0.64 0.92 0.75 0.53 0.82 0.25 F. linae ♂ (N=5) M1 3.79 2.62 0.71 1.27 1.03 0.57 0.86 0.26 M2 3.90 2.84 0.73 1.24 1.08 0.57 0.87 0.27 M3 3.67 2.53 0.65 1.14 0.89 0.53 0.80 0.25 M4 3.67 2.75 0.67 1.21 0.91 0.51 0.86 0.24 M5 3.71 2.67 0.71 1.18 1.03 0.56 0.87 0.25 ♀ (N=5) F1 3.82 2.64 0.69 1.27 0.95 0.61 0.92 0.29 F2 4.07 2.80 0.72 1.22 0.93 0.64 1.02 0.28 F3 3.99 2.80 0.73 1.20 0.93 0.62 0.90 0.30 F4 3.83 2.98 0.82 1.27 0.96 0.69 0.96 0.33 F5 4.17 2.73 0.79 1.18 1.01 0.57 1.03 0.28 F. lindbergae ♂ (N=5) M1 3.47 2.39 0.66 0.89 0.72 0.62 0.90 0.28 M2 3.54 2.51 0.74 0.99 0.72 0.68 0.99 0.29 M3 3.61 2.55 0.78 1.01 0.76 0.65 1.00 0.31 M4 3.40 2.31 0.68 0.95 0.64 0.62 0.95 0.29 M5 4.12 2.85 0.83 1.08 0.71 0.66 1.08 0.29 ♀ (N=5) F1 3.69 2.52 0.76 0.97 0.73 0.62 0.90 0.28 F2 4.24 2.96 0.89 1.05 0.83 0.68 0.99 0.29 F3 4.05 2.83 0.83 0.95 0.72 0.65 1.00 0.31 F4 3.99 2.77 0.84 - 0.71 0.62 0.95 0.29 F5 4.41 3.06 0.92 1.20 0.78 0.66 1.04 0.29 F. liui ♂ (N=3) M1 3.09 2.24 0.62 1.01 0.66 0.55 0.72 0.26 M2 3.40 2.62 0.65 1.00 0.78 0.56 0.71 0.26 M3 3.30 2.34 0.65 1.04 0.71 0.64 0.75 0.30 ♀ (N=3) F1 3.31 2.44 0.68 0.96 0.77 0.56 0.80 0.27 F2 3.42 2.50 0.70 1.04 0.77 0.57 0.80 0.28 F3 3.35 2.46 0.71 0.96 0.69 0.53 0.76 0.25 F. longiceps ♂ (N=5) M1 3.42 2.18 0.65 0.67 0.49 0.42 0.81 0.21 M2 3.13 2.10 0.58 - 0.43 0.45 0.77 0.22 M3 3.10 2.20 0.63 0.65 0.51 0.41 0.79 0.22 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 13
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi M4 3.00 2.11 0.59 0.73 0.50 0.43 0.76 0.22 M5 3.25 2.28 0.65 0.69 0.53 0.41 0.83 0.23 ♀ (N=4) F1 3.48 2.39 0.66 0.65 0.54 0.46 0.92 0.23 F2 3.58 2.44 0.74 0.60 0.49 0.44 0.89 0.22 F3 3.81 2.64 0.77 0.80 0.59 0.50 0.97 0.24 F4 3.44 2.37 0.69 0.60 0.57 0.49 0.90 0.26 M1 3.42 2.18 0.65 0.67 0.49 0.42 0.81 0.21 F. lordhowensis ♂ (N=5) M1 - 2.59 0.68 1.08 0.89 0.47 0.84 0.24 M2 3.84 2.74 0.72 1.11 0.90 0.52 0.81 0.26 M3 3.95 2.79 0.69 1.22 0.94 0.58 0.86 0.31 M4 3.54 2.55 0.64 0.97 0.78 0.48 0.77 0.26 M5 - 2.70 0.71 1.15 0.94 0.53 0.89 0.27 ♀ (N=5) F1 3.93 2.72 0.73 1.04 0.98 0.47 0.86 0.23 F2 4.08 2.88 0.72 1.13 0.96 0.57 0.87 0.27 F3 3.87 2.84 0.74 1.17 0.92 0.58 0.91 0.28 F4 4.46 3.23 0.76 1.13 0.88 0.51 0.92 0.26 F5 4.14 2.91 0.74 1.28 1.00 0.56 0.88 0.29 F. luzonus ♂ (N=1) M1 3.79 2.50 0.67 0.86 0.71 0.51 0.94 0.22 ♀ (N=1) F1 4.07 2.73 0.82 - - 0.52 0.98 0.21 F. madagascariensis ♀ (N=1) F1 4.02 2.76 0.70 - 0.94 0.56 0.96 0.29 F. magnificus ♂ (N=5) M1 3.46 3.46 0.74 0.70 0.60 0.49 0.49 0.24 M2 3.42 3.42 0.68 0.66 0.53 0.46 0.46 0.24 M3 3.55 3.55 0.63 0.72 0.52 0.46 0.46 0.24 M4 3.96 3.96 0.81 - - 0.52 0.52 0.28 M5 3.79 3.79 0.79 0.67 0.51 0.48 0.48 0.24 ♀ (N=5) F1 3.89 3.89 0.75 0.74 0.59 0.49 0.49 0.24 F2 3.92 3.92 0.80 0.78 0.60 0.49 0.49 0.24 F3 4.03 4.03 0.77 0.77 0.59 0.49 0.49 0.25 F4 3.71 3.71 0.66 0.69 0.51 0.44 0.44 0.23 F5 3.62 3.62 0.69 0.77 0.52 0.47 0.47 0.24 F. malayensis ♂ (N=1) M1 3.47 2.71 0.67 - 0.93 0.54 0.83 0.25 ♀ (N=1) F1 3.75 2.77 0.66 1.11 0.93 0.52 0.81 0.25 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F. meilingae ♂ (N=5) M1 4.10 3.03 0.78 1.28 1.06 0.64 0.98 0.30 M2 4.24 2.90 0.79 1.20 0.92 0.55 0.94 0.24 M3 4.02 2.79 0.77 1.28 1.06 0.65 0.89 0.27 M4 4.21 2.88 0.77 1.21 0.98 0.57 0.94 0.27 M5 4.10 2.86 0.74 1.09 0.99 0.59 0.96 0.24 ♀ (N=5) F1 4.42 3.16 0.82 1.07 0.92 0.59 1.02 0.26 F2 4.33 3.03 0.79 1.14 1.06 0.65 1.03 0.29 F3 4.36 2.96 0.79 1.13 0.96 0.60 1.01 0.27 F4 4.27 2.98 0.78 1.14 0.88 0.63 0.97 0.31 F5 4.15 2.91 0.84 1.14 1.03 0.59 0.98 0.27 F. minutus ♂ (N=1) M1 2.61 2.20 0.52 0.85 0.70 0.43 0.61 0.17 F. myersi ♂ (N=5) M1 3.43 2.38 0.57 1.01 0.93 0.52 0.75 0.24 M2 4.18 2.92 0.77 1.37 1.20 0.60 0.94 0.28 M3 3.84 2.78 0.70 1.23 1.08 0.55 0.84 0.27 M4 3.53 2.34 0.66 1.07 1.01 0.55 0.79 0.25 M5 3.38 2.46 0.64 1.05 0.92 0.51 0.76 0.25 ♀ (N=5) F1 4.23 2.90 0.76 1.12 1.08 0.54 0.92 0.27 F2 3.94 2.74 0.74 1.19 0.93 0.57 0.91 0.28 F3 3.73 2.62 0.65 1.04 0.87 0.53 0.80 0.26 F4 4.15 2.84 0.76 1.21 1.07 0.60 0.96 0.29 F5 3.96 2.84 0.68 1.21 1.12 0.57 0.88 0.28 F. nigrescens ♂ (N=3) M1 4.58 3.04 0.73 1.10 0.93 0.70 0.97 0.29 M2 4.17 2.95 0.74 1.06 0.85 0.55 0.93 0.26 M3 4.21 2.83 0.68 1.00 0.89 0.57 0.97 0.25 ♀ (N=3) F1 4.84 3.18 0.79 1.12 0.86 0.58 0.99 0.30 F2 4.57 3.01 0.76 1.02 0.88 0.59 1.02 0.29 F3 4.40 2.84 0.76 1.04 0.83 0.60 1.02 0.30 F. ochraceus ♂ (N=4) M1 3.99 2.92 0.67 - 0.87 0.59 0.27 0.27 M2 3.80 2.71 0.67 1.24 0.92 0.62 0.27 0.27 M3 3.85 2.52 0.67 1.22 0.92 0.61 0.26 0.26 M4 3.56 2.53 0.69 1.17 0.86 0.60 0.25 0.25 ♀ (N=2) F1 3.91 2.71 0.71 - 0.57 0.26 0.26 F2 3.75 2.70 0.71 1.19 0.73 0.63 0.27 0.27 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 15
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F. ovalau ♂ (N=1) M1 3.17 2.83 0.63 0.87 0.65 0.48 0.70 0.24 F. philippinensis ♂ (N=5) M1 3.97 2.77 0.78 1.10 0.77 0.60 0.86 0.24 M2 3.92 2.68 0.77 1.15 0.79 0.56 0.89 0.23 M3 4.03 2.86 0.86 1.09 0.77 0.55 0.91 0.23 M4 4.12 2.84 0.75 1.08 0.79 0.50 0.93 0.23 M5 3.86 2.69 0.74 1.03 0.77 0.53 0.85 0.23 ♀ (N=5) F1 4.30 3.02 0.86 1.03 0.79 0.53 0.95 0.22 F2 4.74 3.20 0.92 1.09 0.85 0.56 1.01 0.24 F3 4.89 3.33 0.93 1.13 0.87 0.61 1.08 0.26 F4 4.65 3.32 0.98 1.24 0.77 0.59 1.04 0.26 F5 4.94 3.42 0.99 1.19 0.85 0.58 1.10 0.26 F. ponaponensis ♂ (N=2) M1 3.50 2.43 0.93 1.00 0.78 0.58 1.00 0.25 M2 3.60 2.55 0.93 1.05 0.85 0.58 0.98 0.28 ♀ (N=2) F1 3.60 2.55 0.93 0.93 0.83 0.55 1.03 0.33 F2 3.80 2.90 1.08 0.98 0.90 0.60 1.13 0.35 F. rubricuneus ♂ (N=2) M1 3.80 2.78 0.75 1.05 0.78 0.53 0.88 0.28 M2 3.60 2.68 0.73 1.03 0.73 0.50 0.80 0.25 F. schuhi ♂ (N=5) M1 3.93 2.68 0.72 1.31 0.97 0.63 0.93 0.29 M2 - 2.76 0.66 1.29 1.06 0.64 0.95 0.31 M3 4.05 2.88 0.76 1.27 0.97 0.62 0.90 0.27 M4 4.18 2.93 0.78 1.40 1.10 0.64 0.96 0.30 M5 4.26 2.89 0.74 1.32 1.08 0.66 0.99 0.30 ♀ (N=5) F1 4.14 2.91 0.74 1.25 0.97 0.62 0.93 0.31 F2 4.35 3.08 0.88 1.43 1.03 0.64 1.04 0.29 F3 4.58 3.20 0.88 1.36 1.03 0.66 1.02 0.29 F4 4.48 3.07 0.79 1.44 1.01 0.66 1.08 0.34 F5 4.53 3.15 0.84 1.40 1.08 0.66 1.08 0.30 F. senaru ♂ (N=3) M1 3.56 2.55 0.69 1.17 0.94 0.51 0.79 0.22 M2 3.83 2.74 0.69 1.08 0.90 0.51 0.81 0.22 M3 3.68 2.60 0.69 1.14 0.88 0.54 0.83 0.25 ♀ (N=3) F1 3.84 2.69 0.70 1.08 0.87 0.54 0.87 0.25 F2 4.10 2.87 0.74 1.12 0.90 0.54 0.90 0.29 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F3 3.97 2.80 0.74 1.21 0.92 0.52 0.88 0.29 F. signis ♂ (N=1) M1 3.30 2.68 0.68 1.10 0.88 0.55 0.78 0.23 ♀ (N=1) F1 3.50 4.95 1.09 4.01 2.21 0.55 0.83 0.23 F. solomonicus ♂ (N=5) M1 2.63 1.86 0.51 0.67 0.58 0.42 0.57 0.21 M2 2.57 1.82 0.47 0.73 0.60 0.42 0.57 0.21 M3 2.77 1.96 0.53 0.83 0.68 0.51 0.60 0.27 M4 2.76 1.88 0.50 0.77 0.55 0.45 0.59 0.24 M5 2.81 1.98 0.52 0.79 0.62 0.46 0.58 0.23 ♀ (N=2) F1 2.92 2.15 0.57 0.80 0.61 0.45 0.68 0.21 F2 2.87 2.11 0.56 0.81 0.70 0.46 0.63 0.23 F .tanna ♂ (N=5) M1 3.51 2.35 0.60 1.02 0.85 0.52 0.79 0.26 M2 3.50 2.38 0.62 0.99 0.88 0.48 0.80 0.24 M3 3.50 2.39 0.59 1.00 0.72 0.48 0.77 0.21 M4 3.55 2.41 0.62 1.08 0.84 0.49 0.80 0.23 M5 3.41 2.27 0.56 1.04 0.73 0.48 0.76 0.23 ♀ (N=5) F1 3.81 2.62 0.65 0.98 0.84 0.51 0.81 0.26 F2 3.56 2.45 0.63 1.00 0.81 0.48 0.76 0.25 F3 3.52 2.34 0.61 1.03 0.82 0.50 0.79 0.24 F4 3.54 2.42 0.60 1.02 0.89 0.54 0.83 0.28 F5 3.48 2.44 0.64 0.94 0.86 0.49 0.84 0.24 F. vitilevu ♂ (N=5) M1 0.57 0.43 0.65 1.12 2.59 0.57 0.83 0.30 M2 1.28 1.02 0.73 1.14 2.77 0.58 0.81 0.29 M3 5.59 3.93 0.81 1.15 2.92 0.60 0.89 0.31 M4 6.88 4.95 0.66 1.03 2.38 0.51 0.74 0.25 M5 - - 0.78 1.19 3.07 0.58 0.92 0.29 ♀ (N=5) F1 6.04 4.35 0.78 1.04 2.91 0.55 0.93 0.28 F2 0.57 0.43 0.80 1.19 3.08 0.63 0.98 0.32 F3 1.28 1.02 0.78 1.17 2.95 0.58 0.92 0.30 F4 5.59 3.93 0.79 1.17 2.90 0.58 0.94 0.31 F5 6.88 4.95 0.87 1.25 3.15 0.66 1.03 0.32 F. wangae ♂ (N=2) M1 4.56 3.24 1.00 1.27 0.99 0.79 1.19 0.32 M2 4.66 3.22 0.92 1.32 1.12 0.79 1.17 0.33 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 17
TABLE 1 (Continued)
Length Width Species Body Cun-Clyp Pronotum AntSeg2 AntSeg1 Head Pronotum InterOcDi F. yasunagai ♂ (N=5) M1 3.29 2.32 0.73 0.65 0.63 0.45 0.83 0.23 M2 3.49 2.54 0.79 0.67 0.67 0.45 0.83 0.21 M3 3.46 2.49 0.77 0.69 0.71 0.46 0.88 0.22 M4 3.43 2.36 0.78 0.67 0.66 0.47 0.84 0.24 M5 3.44 2.41 0.75 0.64 0.57 0.45 0.85 0.23 ♀ (N=5) F1 3.83 2.71 0.85 0.68 0.58 0.48 0.93 0.24 F2 3.77 2.80 0.85 0.60 0.62 0.45 0.99 0.20 F3 3.45 2.65 0.82 0.76 0.68 0.51 0.92 0.24 F4 3.46 2.47 0.73 0.67 0.61 0.49 0.89 0.23 F5 3.76 2.71 0.82 3.67 1.97 1.06 1.21 0.50 F. zuparkoi ♂ (N=5) M1 3.21 2.22 0.69 0.77 0.55 0.47 0.77 0.21 M2 3.16 2.23 0.63 0.72 0.55 0.45 0.81 0.20 M3 3.10 2.20 0.69 0.78 0.56 0.46 0.78 0.21 M4 2.97 2.08 0.67 - - 0.47 0.77 0.22 M5 3.05 2.14 0.69 - 0.48 0.43 0.79 0.19 ♀ (N=5) F1 3.36 2.29 0.70 0.73 0.51 0.48 0.85 0.24 F2 3.33 2.35 0.63 - 0.47 0.43 0.85 0.23 F3 3.38 2.35 0.77 - - 0.52 0.93 0.26 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
Discussion of Major Nodes C-shaped marking on the hemelytron, the elon- gate sclerotized part of ductus seminis and the Nodes of major clades are numbered from 1 absence of vesical spicules, except for F. zuparkoi, to 30. Relationships of Felisacus with other which has a small claw-shaped spicule. Bryocorinae grouping are discussed in Namya- This clade includes F. philippinensis Hsiao, tova et al. (2016) and Namyatova and Cassis 1944, F. longiceps Poppius, 1915, and F. magnifi- (2016). Character-state distributions for each cus (Distant, 1904), which were assigned to the node are as follows: “magnificus group” by Woodward (1954). He Node 1. Felisacus. The genus is supported by also included F. crassicornis Usinger, 1946, and F. five synapomorphies: buccula posterior closed, usingeri Woodward, 1954, but we excluded them binding labial segment I (Namyatova et al., 2016: from the phylogenetic analysis (see Methods and fig. 6D, E) (4-1); labial segment I shorter than or as long as wide (6-0); setae on hemelytron absent Materials). According to Woodward (1954: 49): or present only anteriorly (16-0); parempodia This group is characterised by the subbasally swollen asymmetrical (Namyatova et al., 2016: fig. 20A, first antennal segment, by the nearly straight, scarcely C) (24-1); right paramere longer than left one emarginate base of the pronotum, and by the necklike (figs. 11–13) (29-0). Although these states are postocular part of the head gradually narrowing to unique for Felisacus in this analysis, they also the base, without a cylindrical basal collum sharply occur in other bryocorine groups (see Namya- and completely constricted from a wider region tova et al., 2016). Felisacus is characterized by the behind the eyes. In magnificus, philippinensis, and unique combination of characters of the thorax, crassicornis, at least, the right clasper [= paramere] of labium, pretarsus, and male genitalia (see generic the male has the form of a curved rod, slender diagnosis below). throughout its length, not considerably widened and Node 2. Felisacus curvatus Hu and Zheng, flattened basally; the left clasper is nearly straight and 2001, + F. wangae, sp. nov. The node is supported only feebly curved on its outer margin; the right by two contradicted synapomorphies: labial seg- clasper is slightly knobbed or spatulate at apex, and ment I twice as long as wide (6-1); dorsal labiate either the right or the left clasper, or both, may give plate distinctly striated (fig. 16L, N) (70-1). F. off a shorter or longer internal branch; ventral margin curvatus and F. wangae are very distinct species of apex of abdomen narrowly convex between right not very similar to each other or any other Feli- and left claspers. sacus species. Node 3. This clade is supported by four We did not find that any of the characters uncontradicted synapomophies: left paramere described by Woodward (1954) delimit this with clump of setae on inner angle (51-1) (e.g., group and were not recovered as figs. 11T, AB, 12M, W); at least apical half of synapomorphies. ductus seminis sclerotized (52-2) (e.g., fig. 10A, Node 4. Felisacus luzonus, sp. nov., and F. C, D); dorsal labiate plate very small and thin, philippinensis. This clade is supported by three placed entirely between valvulae apodemes (fig. contradicted synapomorphies: vertex flat (as in 16E, G, K) (67-1); spermathecal gland placed at Namyatova et al., 2016: fig. 6E) (0-0); distance the anterior margin of dorsal labiate plate (e.g., between transverse depression and pronotum as fig. 16E, G) (71-1). It is also supported by a con- long as or longer than eye diameter (3-2); apical tradicted synapomorphy: distinct teeth on vesica part of right paramere distinct (31-1). Felisacus and spicules absent (66-0). All species in this luzonus and F. philippinensis are also restricted to clade also have the inner margin of the cuneus the Philippines. straight (as in Namyatova et al., 2016: fig. 13D), Node 5. This node is supported by two syn- that is unique within the genus. Many species apomorphies: inner margin of middle part of within this clade are characterized by the right paramere without swelling, straight or con- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 19
Stenotus binotatus Nesidiocoris tenuis Pachypeltis reuteri 1 Felisacus curvatus 2 96 Felisacus wangae Felisacus bismarckensis Felisacus ponaponensis 1 Felisacus luzonus 4 99 1 95 Felisacus philippinensis 3 97 Felisacus ceylonicus Felisacus cristobalus Felisacus fedori 1 Felisacus lindbergae 8 7 1 Felisacus longiceps 97 95 Felisacus magnificus Felisacus yasunagai Felisacus zuparkoi Felisacus bradi Felisacus ovalau Felisacus vitilevu Felisacus webbi Felisacus bau 2 14 Felisacus bellus 96 Felisacus caledonicus Felisacus dauloi Felisacus elegantulus Felisacus filicicola 1 17 Felisacus jacobsoni 96 Felisacus javanus Felisacus lambkinae Felisacus liui Felisacus lordhowensis Felisacus malayensis Felisacus minutus Felisacus nigrescens 18 Felisacus rubricuneus Felisacus senaru Felisacus schuhi Felisacus signis Felisacus solomonicus Felisacus tanna 2 Felisacus australicus 24 97 Felisacus ochraceus Felisacus amboinae Felisacus linae 4 Felisacus meilingae 26 99 Felisacus myersi Felisacus albus 1 Felisacus glabratus 30 95 Felisacus indicus Felisacus insularis
FIGURE 1. Strict consensus tree, unweighted analysis. The numbers above branches correspond to the Bremer supports values and those below branches correspond to the Bootstrap support values. 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 cave (e.g., fig. 11R, T, AB) (34-0); outgrowth on Node 11. This node is supported by a single posterior side of left paramere apically present noncontradicted synapomorphy: elongate out- (e.g., figs. 12N, R, 13J, AC) (43-1). State (34-0) is growth on inner angle of right paramere straight contradicted. Felisacus ponaponensis (Carvalho, (figs. 11AB, 12M) (40-0). 1956) has a basal position and differs from other Node 12. This node is supported by a single species of this clade in many characters (see dis- contradicted synapomorphy: outer margin of cussion for this species below). left paramere straight or convex (figs. 12N, Node 6. This node is supported by three con- 13AC) (46-0). tradicted synapomorphies: posterior part of pro- Node 13. Felisacus cristobalus, sp. nov., + F. notum uniformly brown, dark brown to black fedori, sp. nov. This node is supported by two (14-2); C-shaped marking on corium present (21- contradicted synapomorphies: dorsal side of 1); middle part of right paramere narrower than head posteriorly at least slightly swollen (as in basal part (e.g., figs. 11L, T, 12M, U) (35-2). This Namyatova et al., 2016: fig. 6D) (0-1); distance node comprises all species with both arms of between transverse depression on head and pro- C-shaped marking on the hemelytron developed. notum shorter than eye diameter (3-1). Also, it Node 7. This node is supported by five syn- is supported by the noncontradicted synapomor- apomorphies: dorsal side of head flat (as in phy—the C-shaped marking on the hemelytron, Namyatova et al., 2016: fig. 6E) (0-0); distance with anterior arm short, not reaching R+M (22- between transverse depression and pronotum as 1). F. cristobalus and F. fedori are very similar to long as or longer than eye diameter (3-2); swell- each other in male genitalic structure, as well as ing or outgrowth at the inner margin of proximal the C-shaped marking on the hemelytron. inflection of right paramere present (e.g., fig. Node 14. This node is supported by two non- 12M, Q, W) (38-1); outgrowth on inner margin contradicted synapomorphies: dorsal surface of of left paramere elongate (figs. 11S, 12R, X) (48- labial segment II elongate, covering base of seg- 2); distal sclerotized part of ductus seminis ment III (8-1) (Namyatova et al., 2016: fig. 6D); curved (e.g., fig. 10C, D, F) (53-1). Only state semioval sclerite on dorsal labiate plate present 38-1 is noncontradicted. Felisacus ceylonicus, sp. (fig. 16C; as in Namyatova et al., 2016: fig. 23F) nov., has a basal position in this clade, but is (68-1). It is also supported by two contradicted similar to F. magnificus Distant, 1904, F. lindber- synapomorphies: labial segment III subequal to gae, sp. nov., and F. longiceps Poppius, 1915. or longer than ventral side of head (Namyatova Node 8. This node is supported by a single et al., 2016: fig. 6E) (9-1); outer margin of right noncontradicted synapomorphy: small tooth on paramere straight, slightly concave or convex posterior margin of genital capsule present (e.g., (e.g., fig. 11A, C, H) (3-0). The state 9-1 is unique fig. 14I, T, V) (26-1). F. magnificus has a basal within Felisacus. The species within this clade position in this clade. Externally it is almost mostly have a pale hemelytron and are very simi- indistinguishable from F. longiceps. lar. However, three basal species, F. bradi, sp. Node 9. This node is supported by a single syn- nov., F. ovalau, sp. nov., and F. vitilevu, sp. nov., apomorphy: outer margin of left paramere dis- differ from the rest by the structure of the sec- tinctly concave (figs. 11U, AC, 12R, 13AE) (46-1). ondary gonopore, which is more or less enlarged Node 10. This node is supported by a single or elongate (figs. 9N, 10E, M), whereas it is small contradicted synapomorphy: outgrowth on dor- and cuplike in the remaining representatives of sal side of apical part of left paramere present this clade (e.g., fig. 8A–D). (e.g., figs. 11C, AC, 12N, R, 13AC) (50-1). This Node 15. This node is supported by a single character also present in Felisacus ceylonicus, sp. noncontradicted synapomorphy: transverse nov., which is similar externally to F. longiceps depression delimiting occipital region distinct, and F. lindbergae. surrounding neck (Namyatova et al., 2016: figs. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 21
Stenotus binotatus Nesidiocoris tenuis Pachypeltis reuteri Felisacus curvatus 2 Felisacus wangae Felisacus luzonus 4 Felisacus philippinensis 3 Felisacus ponaponensis 5 Felisacus bismarckensis 6 Felisacus ceylonicus 7 Felisacus magnificus 1 8 Felisacus zuparkoi 9 Felisacus longiceps Felisacus lindbergae 10 12 Felisacus yasunagai 11 Felisacus cristobalus 13 Felisacus fedori Felisacus bradi Felisacus ovalau 14 16 Felisacus vitilevu 15 Felisacus webbi Felisacus rubricuneus 17 Felisacus bau Felisacus senaru 18 21 Felisacus lambkinae 22 Felisacus javanus 23 19 24 Felisacus australicus Felisacus ochraceus Felisacus amboinae Felisacus bellus Felisacus caledonicus 20 Felisacus dauloi Felisacus elegantulus Felisacus jacobsoni Felisacus linae Felisacus liui Felisacus malayensis Felisacus schuhi 25 Felisacus signis Felisacus meilingae 26 Felisacus myersi Felisacus minutus 27 28 Felisacus nigrescens Felisacus solomonicus Felisacus filicicola 29 Felisacus lordhowensis Felisacus tanna Felisacus albus Felisacus glabratus 30 Felisacus indicus Felisacus insularis
FIGURE 2. Strict consensus tree, implied weight analysis, k = 7. 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
4E, 6D) (1-2). The node includes all species of and narrowed and curved apically, the inner margin node 11 except for F. bradi. Externally this spe- with a more or less pronounced subbasal expansion; cies is similar to F. ovalau. the left clasper is strongly curved upward and Node 16. The node is supported by two con- inward, with its outer margin strongly convex; ven- tradicted synapomorphies: apex of genital cap- tral margin of apex of abdomen widely convex sule curved dorsally (fig. 15F, L) (27-1); inner between right and left claspers. margin of middle part of right paramere without The cylindrical antennal segment I and con- swelling, straight or concave (34-0) (fig. 13E, V). striction of the postocular region are contra- This clade includes F. ovalau and F. vitilevu. dicted synapomorphies. The species of this group Although the clade has insignificant resampling also have very similar or identical parameres. support, both species are similar to each other Node 18. This node is supported by two syn- externally and in genital morphology, and both apomorphies: inner part of clavus dark brown to are known from Fiji. black (18-1); tooth on posterior side of left para- Node 17. The node is supported by three mere medially present (fig. 11G) (44-1). The for- contradicted synapomorphies: transverse mer is contradicted and the latter is depression surrounding occipital region (as in uncontradicted. The species of this clade are Namyatova et al., 2016: fig. 4E) (2-2); antennal similar to each other externally, varying mostly segment I straight (as in Namyatova et al., 2016: in coloration. They also have similar structure of fig. 8A) (5-0); distinct teeth or spicules on the male genitalia, including genital capsule, vesica absent (66-0). State 5-0 is unique in Feli- parameres, and vesica with elongate spicules. The sacus; F. webbi, sp. nov., the only species known species of this node mostly differ in shape and from Madagascar included in the analysis, is number of those spicules. sister to the remainder of this clade, and differs Node 19. This node is supported by two from other species of this group by having only uncontradicted apomorphies: spicule A (convo- a single curved vesical spicule (fig. 10N). In luted, placed on right-hand side) present (e.g., contrast, other species have many vesical spic- fig. 8A–E) (54-1); spicule C (convoluted, placed ules. Although the clade is relatively well sup- on left-hand side) present (e.g., fig. 8B, D, N) ported, the relationships of F. webbi with other (58-1). This clade includes all species from clade species are doubtful. 18 except for F. rubricuneus Carvalho, 1956. This node can be associated with the Wood- Node 20. This node is supported by two syn- ward’s (1954) “elegantulus” group, as the latter apomorphies: apical part of right paramere more includes F. elegantulus (Reuter, 1904), F. filicicola or less distinctly demarcated (e.g., fig. 11C, E, J, (Kirkaldy, 1908), F. jacobsoni Poppius, 1914, F. P) (31-1); spicule B (curved, placed from right- ochraceus Usinger, 1946, and F. amboinae Wood- hand side (e.g., fig. 8B, E, F). This clade includes ward, 1954. He also included F. nigricornis Pop- all species from clade 18 except for F. bau, sp. pius, 1912, which was excluded from our analysis nov., and F. rubricuneus. (see Methods and Materials). Woodward (1954: Node 21. This node is supported by a single 49) defined this group as having uncontradicted synapomorphy: outer margin of cylindrical first antennal segment, not swollen sub- right paramere distinctly concave (figs. 11E, 12G, basally, the distinctly excavated base of the prono- I, 13D, N) (33-1). This clade comprises F. austra- tum, and the differentiation of the base of the head licus, sp. nov., F. javanus Reuter, 1908, F. lambki- as a cylindrical collum sharply and completely con- nae, sp. nov., F. ochraceus Usinger, 1946, F. stricted from a wider region just behind the eyes. In senaru, sp. nov. These species also possess similar F. elegantulus F. filicicola, F. ochraceus, and F. amboi- vesical spicules (figs. 8C, N, O, 9F, I). nae, at least, the right clasper [ = paramere] of the Node 22. This node is supported by a single male is considerably widened and flattened basally contradicted synapomorphy: base of spicule C 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 23
Stenotus binotatus Nesidiocoris tenuis 66 0 Pachypeltis reuteri 0 3 21 6 70 Felisacus curvatus 2 0 2 1 1 1 Felisacus wangae 37 41 21 27 33 0 3 31 1 1 0 Felisacus luzonus 1 1 4 6 32 53 0 2 1 Felisacus philippinensis 0 1 5 10 11 12 13 15 17 23 51 52 66 67 71 2 18 48 1 2 0 1 1 3 1 1 1 1 1 1 1 1 1 1 2 0 1 Felisacus ponaponensis 34 43 0 1 5 Felisacus bismarckensis 14 21 35 50 6 1 Felisacus ceylonicus 2 1 2 45 0 3 38 48 53 7 Felisacus magnificus 0 2 1 2 1 1 49 26 Felisacus zuparkoi 4 6 16 24 29 1 8 1 46 1 0 0 1 0 1 Felisacus longiceps 1 9 48 50 46 0 Felisacus lindbergae 1 10 18 34 35 39 0 12 40 0 1 0 0 Felisacus yasunagai 11 39 49 0 0 3 22 0 1 Felisacus cristobalus 13 25 28 31 1 1 1 Felisacus fedori 70
1 Felisacus bradi 14 47 8 9 33 68 27 34 2 2 Felisacus ovalau 33 35 36 37 41 1 1 0 1 14 1 0 16 2 2 0 1 1 Felisacus vitilevu 1 Felisacus webbi 2 15
2 5 66 Felisacus rubricuneus 14 2 0 0 17 Felisacus bau 1 14 18 44 18 1 Felisacus senaru 1 1 33 1 21 Felisacus lambkinae 59 22 54 58 0 18 Felisacus javanus 1 1 19 0 23 57 65 Felisacus australicus 24 1 1 Felisacus ochraceus Felisacus amboinae 31 56 20 Felisacus linae 1 1 14 18 1 1 Felisacus malayensis
59 Felisacus schuhi 0 55 60 62 64 Felisacus meilingae 1 1 1 1 26 56 Felisacus myersi 35 18 32 0 34 58 1 Felisacus minutus 25 14 18 0 0 27 0 0 30 Felisacus nigrescens 28 2 1 1 Felisacus solomonicus 57
1 Felisacus caledonicus Felisacus dauloi 64 14 18 Felisacus bellus 1 1 1 Felisacus jacobsoni 58
0 Felisacus elegantulus 14 33 1 1 Felisacus liui 18 1 Felisacus signis
54 61 Felisacus lordhowensis 64 0 29 54 1 1 Felisacus filicicola 35 1 1 Felisacus tanna Felisacus albus 33 35 63 Felisacus glabratus 30 1 1 1 Felisacus indicus Felisacus insularis
FIGURE 3. One of the shortest trees found in the analysis using implied weights, K = 7. The white circle cor- responds to homoplasy, the characters occurring in two or more clades. The black circle corresponds to apomorphy, the character unique for the clade, even if it is lost in some its members. The characters were mapped using unambiguous optimization. 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
(convolute, placed from left-hand side) approxi- fer slightly from each other in vesical spicules mately on the same level with other spicules and coloration (see discussion for those species), (figs. 8C, N, O, 9F) (59-0). The clade includes overall they are very similar to each other exter- four species: F. australicus, sp. nov., F. javanus nally and in genital monophyly, and they are Reuter, 1908, F. lambkinae, sp. nov., and F. ochra- both distributed in Vanuatu. ceus Usinger, 1946. All those species have similar Node 27. This node is supported by two coloration with a red cuneus and they also have contradicted apomorphies: inner margin of similar parameres and vesical spicules (figs. 8C, middle part of right paramere without swell- N, O, 9F). Felisacus javanus and F. lambkinae ing, straight or concave (figs. 12AC, 13A, R) have almost identical spicules (fig. 8N, O), and (34-0); spicule C (convolute, placed from left- the spicules in F. australicus and F. ochraceus are hand side) absent (fig. 9D, E, L) (58-0). This also very similar (figs. 8C, 9F). clade includes F. minutus Carvalho, 1981, F. Node 23. This node is supported by a single nigrescens Carvalho, 1981, and F. solomonicus, uncontradicted synapomorphy: inner part of cla- sp. nov., which are not similar to each other vus transparent, whitish to pale brown (18-0). externally, and the proposed close relationship This clade comprises F. javanus, F. australicus, in this analysis is doubtful. and F. ochraceus. Node 28. This node is supported by a single Node 24. This node is supported by two syn- uncontradicted synapomorphy: apex of right apomorphies: apical arm of spicule B (curved, paramere distinctly concave (fig. 13A, R) (30-1). placed from right-hand side) (figs. 8C, 9F) (57- This clade comprises F. nigrescens and F. solo- 1); spicule H (hook shaped, placed from left- monicus. Although those two species are very hand side) present (figs. 8C, 9F) (65-1). This different externally and their vesical spicules are clade includes F. australicus and F. ochraceus. not very similar (fig. 9E, L), they have strikingly Those two species are very similar to each other similar right parameres and they are geographi- structurally and have nearly identical male gen- cally proximal, with F. nigrescens known from italia (figs. 8N, 9F), but they differ in coloration New Guinea and F. solomonicus known from (see diagnoses and discussions for these spe- Solomon Islands. cies). They are widely separated species geo- Node 29. The node is supported by a single graphically, with F. ochraceus from Guam contradicted synapomorphy: spicules E (weekly (Mariana Is.) and F. australicus from Queensland sclerotized, placed from right-hand side) present (Australia). (figs. 8I, R, 9M) (61-1). The clade includes F. Node 25. This node is supported by two non- filicicola (Kirkaldy, 1908), F. lordhowensis, sp. contradicted synapomorphies: inner part of cla- nov., and F. tanna, sp. nov. These three species vus transparent, whitish to pale brown (18-0); are similar externally, including pale coloration. apical part of right paramere as long as or longer Node 30. This node is supported by two syn- than medial part (e.g., fig. 11C, J, P) (32-0). apomorphies: middle part of right paramere Node 26. F. myersi, sp. nov., + F. meilingae, sp. twice as wide as basal part (figs. 11A, AF, 12A, nov. This node is supported by three uncontra- C) (35-1); spicule F (long, needle shaped, placed dicted apomorphies: spicule A (convoluted on from left-hand side), present (fig. 8A, J, K, L) right had side) almost straight (55-1); spicule D (63-1). The former state is contradicted and the (small, placed basally) present (60-1); spicule I latter is uncontradicted. This clade comprises (convolute basally, medial) present (62-1); and four species distributed from the Oriental region: by a single contradicted synapomorphy: spicules F. albus, sp. nov., F. glabratus (Motschulsky, G (long and distinctly concave apically, arising 1863), F. indicus, sp. nov., and F. insularis Miya- from left-hand side) present (fig. 9B, C) (64-1). moto, 1965. The species of this clade are similar Although, species F. myersi and F. meilingae dif- in the structure of the spicules and parameres, as 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 25 well as external morphology and pale riorly to sulcus (Namyatova et al., 2016: fig. 4E); coloration. unique structure of pleura with distinct triangu- lar scent gland evaporative area, oval spiracle, surrounded with few evaporative bodies; meso- Taxonomy pleural/metapleural suture inferiorly obsolete Felisacus Distant, 1904 (Namyatova et al., 2016: fig. 14B); parempodia asymmetrical, outer parempodium shorter than Liocoris Motschulsky, 1863: 86 (gen. nov.; type inner one (Namyatova et al., 2016: fig. 20A, C). species Liocoris glabratus Motschulsky, 1863, Description: Male. Body small and elongate, by monotypy); Distant, 1904: 438 (junior 2–4 mm. COLORATION (figs. 4–7): Head, pro- homonym of Liocoris Fieber, 1858 (Hemip- notum, and thoracic pleura usually yellow or tera); Carvalho, 1957: 103 (catalog); Schuh, pale brown, sometimes with brown to dark 1995: 510 (catalog). brown marking or uniformly dark brown to Felisacus Distant, 1904: 438 (replacement name black; hemelytron often whitish yellow and for Liocoris Motschulsky, 1863); Kirkaldy, translucent, often with brown to dark brown 1906: 134 (list); Reuter, 1910: 163 (catalog); C-shaped marking or at least with dark stripe Poppius, 1911: 3 (description); Poppius, along inner margin of corium; coloration of 1912: 181 (description); Bergroth, 1922: 55 cuneus varying from whitish yellow to dark (list); China, 1944: 174 (key to genera); Car- brown, sometimes with reddish tinge or bright valho, 1952: 55 (catalog); Woodward, 1954: red; membrane sometimes with grayish tinge or 41 (discussion, description); Carvalho, 1955: brown; appendages of same color or paler than dorsum, usually whitish yellow, yellow or pale 34 (key to genera); Carvalho, 1957: 103 (cat- brown, antennal segment I sometimes dark alog); Woodward, 1958: 236 (discussion); brown to black; femur sometimes with brown or Carvalho, 1981: 51, 61 (key to genera, reddish bands. SURFACE AND VESTITURE: description, key to species); Cassis and Body mostly smooth, often with shallow punc- Gross, 1995: 141 (catalog); Schuh, 1995: 510 tures; lateral margins of scutellum with shallow (catalog); Lin, 2000: 233 (key to species). striations (Namyatova et al., 2016: fig. 11A, B); Hyaloscytus Reuter, 1904: 1 (original description; R+M impunctate or with shallow punctures; type species: Hyaloscytus elegantulus Reuter, body clothed with pale simple setae, mostly 1904, by monotypy); Kirkaldy, 1906: 134 missing on hemelytron, only with few setae on (list); Reuter, 1910: 153 (catalog); Poppius, clavus and anterior margin of corium; trichoboth- 1911: 3 (synonymy); Carvalho, 1957: 103 ria surrounded by dense fields of trichoma. (catalog); Schuh, 1995: 510 (catalog) STRUCTURE: Head: Hypognathous, in dorsal Felisacoris Carvalho, 1956: 24 (gen. nov.; type view eyes distinctly removed from pronotum, species: Felisacoris ponaponensis Carvalho, roundish; area between head to pronotum 1956, by monotypy); new synonymy. straight or slightly swollen, sometimes with transverse depression, delimiting occipital Diagnosis: Felisacus is separated from other region; longitudinal sulcus present, its length bryocorine taxa by the following characters: varying from very short to almost reaching ante- labial segments I and II almost as long as wide rior margin of head; frons straight, not convex (Namyatova et al., 2016: figs. 6D, E, 9C ), some- (Namyatova et al., 2016: fig. 4E); in anterior view times twice as long as wide, but each of them head slightly higher than wide, eye oval, shorter distinctly shorter than segment III; pronotum than head height, its inner margin distinctly con- with two pairs of punctures laterally on sulcus cave; clypeus as long as eye, its base placed above delimiting calli and with pair of punctures ante- inferior margin of eye; antennal fossa tubercu- 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 late, shorter than half of eye height, attached near epimeron narrow, without lobe; posterior margin midline of eye, its inferior margin not reaching of metasternum straight. Hemelytron (Namya- inferior margin of eye (Namyatova et al., 2016: tova et al., 2016: figs. 11A, 13E, F): Clavus with fig. 3C); in lateral view eye oval, its inferior mar- row of punctures, medial margin longer than gin not reaching maxillary plate; antennal fossa scutellum; medial fracture very short; R+M long, inserted slightly above base of clypeus; clypeus reaching cuneus, impunctate or with some very slightly swollen basally; buccula short and ring- small and shallow punctures; claval commissure like, tightly binding labial segment I; gula long with straight margins; cuneus 2–4 times as long and straight, 2–3× as long as buccula (Namya- as wide at base, its inner margin straight or con- tova et al., 2016: fig. 6D, E). Antenna: Slightly vex; membrane with single or two cells, often shorter than or subequal to body length; length vein between them faint; cells not reaching apex of segment I slightly greater than or subequal to of membrane, forming right angle, shorter than head width, cylindrical or swollen medially distance between apex of cell and apex of mem- (Namyatova et al., 2016: fig. 8A, B); segment II brane. Legs: Forecoxae contiguous with each subequal to or slightly longer than segment I, other, middle and hind coxae separated (Namya- segment III as long as or slightly longer than seg- tova et al., 2016: fig. 17C), in lateral view coxae ment II; segment IV usually slightly shorter than short, not reaching upper margin of thoracic or subequal to half the length of segment I. pleura; femora slender, not modified (Namyatova Labium (Namyatova et al., 2016: figs. 6D, E, 9C): et al., 2016: fig. 18E), forefemur shorter than Length variable, reaching from posterior margin length of head and pronotum combined; hind of prosternum to posterior margin of metaster- femur longer than other femora; tibia straight; num; labial segments I and II shortened, often tarsal segments almost subequal in length, seg- wider than long or as long as wide, rarely some- ment III dilate (Namyatova et al., 2016: fig. 19C). what longer than wide; labial segment III some- Pretarsus (Namyatova et al., 2016: fig. 20A–C). times telescoped into segment II; segment III Unguitractor with three rows of contiguous tiles, usually 2–3 times longer than, rarely as long as those of middle row straight; basal claw spicules segments I and II combined; segment IV usually absent; claw broadly rounded, often with basal almost twice as long as segment III, rarely only subtriangular tooth, sometimes only with short slightly longer than segment III. Thorax: Prono- outgrowth basally; claw without medial or apical tum (Namyatova et al., 2016: figs. 4E, 6D). tooth; parempodia setiform, tapering toward Length of pronotum subequal to its width at apex, asymmetrical, with outer parempodium base; collar often more or less delimited laterally shorter than inner one; pseudopulvilli present, and medially, in F. ponaponensis collar not delim- attached to unguitractor and base of claw, dis- ited; calli fused, delimited with sulcus posteri- tinctly longer than claw. Genitalia: Genital cap- orly; pair of punctures of lateral side of this sule (figs. 14, 15; Namyatova et al., 2016: fig. 22L) sulcus and anteriorly to sulcus present. Scutel- subtriangular, 1–1.5× as long as wide; without lum and mesoscutum (Namyatova et al., 2016: supragenital bridge, sometimes with outgrowths fig. 11A, B). Sulcus between scutellum and meso- on apical margin of ventral wall and swelling at scutum with pair of punctures medially; apex of sides. Parameres (Namyatova et al., 2016: figs. scutellum acute. Pleura (Namyatova et al., 2016: 11–13, fig. 22J, K). Right paramere longer than fig. 14B). Meso- and metapleuron not separated, left, parameres usually sickle shaped or L-shaped, suture between them obsolete inferiorly; meso- but sometimes right paramere straight or left thoracic apodeme round; metathoracic spiracle paramere of irregular shape. Aedeagus (Namya- open and oval, with single row of evaporative tova et al., 2016: figs. 8–10, figs. 22I, M) phal- bodies inferiorly; opening of scent gland distinct, lotheca moderately sclerotized; endosoma surrounded by triangular evaporative area; met- voluminous, subdivided into vesica and conjunc- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 27 tiva (fig. 10; Namyatova et al., 2016: figs. 22I, M), are distributed in southeast Asia. Only five species with or without long spicules, rarely with fields are known from Australian region, including a of small spicules or with single small claw-shaped single species each from mainland Papua New spicule; ductus seminis with small sclerotized Guinea, New Britain, and Micronesia, and two ring around secondary gonopore, sometimes species from the Solomon Islands (fig. 22). Species apical half of ductus seminis sclerotized, straight of clade 14 are distributed in the Australian and or hook shaped apically. Oriental regions, ranging from southern China Female. Similar to male, not showing sexual and Japan to the southeast coast of Australia and dimorphism in external characters, but females New Zealand; it is also widely distributed in usually slightly larger than males. Genitalia (fig. islands of the southwest Pacific and occurs in 16; Namyatova et al., 2016: figs. 23F, G): Dorsal Madagascar (fig. 22). This pattern is similar to labiate plate with or without sclerotized rings, Coridromius, which is distributed in Indo-Pacific sometimes with semioval sclerite medially; often although its two main subclades are largely without striations, smooth, sometimes with stria- divided at Wallacea (Tatarnic and Cassis, 2010). tions, rarely with membranous longitudinal ridge; Host plants: Little is known about the feed- lateral oviducts attached medially or in posterior ing habits of Felisacini, but they are assumed to half; place of attachment on spermathecal gland be fern feeders based on their common and spe- varying, often medially or near anterior margin; cialized association with ferns (15 species of Feli- posterior wall of bursa copulatrix membranous, sacus of 55 described with host-plant data). with or without small tubercles, sometimes with Fourteen species have been collected from ferns membranous lobe anteriorly; vulva indistinct, (Kirkaldy, 1908; Usinger, 1946; Miyamoto, 1965; without sclerotization. Woodward, 1958; also, this work). The host-plant Distribution: The genus is mostly distrib- records of Felisacus do not indicate phylogenetic uted in tropical areas of the Indo-Pacific region. restrictedness to any fern clade, and are instead Felisacus is distributed mainly in Australian and known from distantly related genera, including Oriental regions, with 34 and 22 species respec- Nephroplepis Schott, Polystichum Roth (Polypo- tively. Three species are also known from the diaceae), Diplazium Schwartz, Plesioneuron Holt- Ethiopian region. Felisacus jacobsoni, F. javanus, tum (Aspleniaceae), Pteris L. (Pteridaceae), F. longiceps, and F. magnificus are distributed in Hypoplepis Bernh., Pteridium Gled. ex Scop. the Australian and Oriental regions. Most spe- (Dennstaedtiaceae), Schizaea Sm. (Schizaeaceae) cies inhabit small areas, especially, those found (see Christenhusz and Chase, 2014, for the latest on islands; however, some species have broader phylogeny of ferns). Moreover, there are cases distribution: F. bellus Lin, 2000 (from southern where a Felisacus species is known from two or China to southern parts of Vietnam and Thai- more species, with these multiple fern species land), F. elegantulus (east coast of Australia and belonging to different families. This broad fern New Zealand), F. filicicola (Samoa, Fiji, Vanuatu), affiliation is also found in the tribe Bryocorini F. insularis (southern China, Japan, Thailand, (Konstantinov and Knyshov, 2015). Felisacus Vietnam), F. javanus (from Malay to New elegantulus, F. lindbergae, and F. nigrescens have Guinea), F. longiceps (from South China to been recorded from angiosperms, but these likely Ambon Is.), F. magnificus (from Thailand and represent sitting records, because records are Japan to Lombok Is.). either singletons or doubletons. The phylogeny of Felisacus is divided into two Discussion: Felisacus is well delimited and main clades (3 and 14) and a single small Asian although the species within it vary considerably in clade 2 with unclear affinities (figs. 1–3). Clades 3 salient and genital structures, the genus is not and 14 are broadly overlapping in the Oriental similar to any other genus of Miridae. Here we region. Clade 3 includes 12 species of which eight synonymize Felisacoris Carvalho, 1952, with Feli- 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 sacus. Although the single species of Felisacoris, F. Carvalho (1957) and Schuh (1995). Under each ponaponensis, is morphologically distinct, it is specimen treatment only citation of the original undoubtedly nested within Felisacus, in particular description and synonymy is given. in clade 3, and possesses male genitalia very simi- Felisacus adamsi Carvalho, 1956 (Micronesia) lar to other species in this group (see figs. 1–3, 10). Felisacus albus, sp. nov. (Thailand) Although Felisacus is a speciose taxon, contain- Felisacus amboinae Woodward, 1954 (Ambon Is.) ing 55 species, there are more undescribed species Felisacus auritulus Distant, 1913 (Seychelles) known to us. We examined such specimens Felisacus australicus, sp. nov. (Australia) belonging to the Bishop Museum, and likely rep- Felisacus bau, sp. nov. (Borneo) resent new species, but we refrained from treating Felisacus bellus Lin, 2000 (South Asia) them as they were in poor condition. Felisacus bismarckensis, sp. nov. (New Britain) Confusion exists in the literature concerning Felisacus bradi, sp. nov. (Society Is.) the type species and synonymy of the valid Felisacus caledonicus, sp. nov. (New Caledonia) genus and junior synonyms. In particular, the Felisacus capitatus Miyamoto, 1965, junior syn type species of Liocoris is L. glabratus, which onym of F. magnificus Carvalho (1957) listed as being synonymous Felisacus carpenterae Hsiao, 1944, junior syn- with Hyaloscytus elegantulus, which he attrib- onym of F. javanus uted to Poppius (1911). This is in error in both Felisacus ceylonicus, sp. nov. (Sri Lanka) the relevant available names and the date of Felisacus crassicornis Usinger, 1946 (Guam) Poppius’ work. Poppius (1911) redescribed Feli- Felisacus cristobalus, sp. nov. (Solomon Is.) sacus and F. glabratus, synonymized Hyaloscytus Felisacus curvatus Hu and Zheng, 2001 (South with Felisacus, but made no mention of species Asia) synonymy. Carvalho (1957) indicated that Pop- Felisacus dauloi Woodward, 1958 (Papua New pius (1911) synonymized Felisacus glabratus Guinea) with Hyaloscytus elegantulus. Kerzhner and Felisacus elegantulus (Reuter, 1904) (Australia) Jansson (1985) indicated that Carvalho’s citation Felisacus fedori, sp. nov. (Sulawesi) of Poppius was incorrect, and identified Poppius Felisacus filicicola (Kirkaldy, 1908) (Fiji, Samoa, (1914) as the work with the synonymy of Felias- Vanuatu) cus glabratus and Hyaloscytus elegantulus java- Felisacus glabratus (Motschulsky, 1863) (Sri nus, and not the nominotypical subspecies of Lanka) the latter. Cassis and Gross (1995) followed the Felisacus gressitti Miyamoto, 1965 (Japan) Poppius attribution of Kerzhner and Jansson Felisacus indicus, sp. nov. (India) (1985) but also indicated that H. elegantulus was Felisacus insularis Miyamoto, 1965 (South Asia) synonymous with F. glabratus (as Liocoris), fol- Felisacus jacobsoni Poppius, 1914, stat. nov. (Java, lowing Carvalho’s (1957) species nomenclatorial Lombok Is.) history of F. glabratus. Schuh (1995) cited Lioco- Felisacus javanus Reuter, 1908 (South Asia) ris glabratus as the type of Felisacus and gave it Felisacus lambkinae, sp. nov. (Christmas Is.) as a separate species with no synonyms. We Felisacus linae, sp. nov. (East Timor) revise the status of Hyaloscytus elegantulus java- Felisacus lindbergae, sp. nov. (Solomon Is.) nus in this work and raise it to species rank. Felisacus liui, sp. nov. (Solomon Is.) Felisacus longiceps Poppius, 1915 (South Asia) Felisacus lordhowensis, sp. nov. (Lord Howe Is.) Checklist of Felisacus species Felisacus luzonus, sp. nov. (Philippines) Valid names are given in italics; synonyms are Felisacus madagascariensis Poppius, 1912 (Mada- given in regular type. Full nomenclatorial listings gascar) for previously described species can be found in Felisacus magnificus Distant, 1904 (South Asia) 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 29
Felisacus malayensis, sp. nov. (Penang, Malaysia) (as in Namyatova et al., 2016: fig. 6D); scler- Felisacus meilingae, sp. nov. (Vanuatu) otized part of ductus seminis straight (fig. Felisacus minutus Carvalho, 1981 (Papua New 10A...... F. bismarckensis, sp. nov. (fig. 4) Guinea) – Distance between longitudinal sulcus of vertex Felisacus myersi, sp. nov. (Vanuatu) and anterior margin of pronotum as long as Felisacus nigrescens Carvalho, 1981 (Papua New eye diameter; vertex flat or only slightly Guinea) upraised (Namyatova et al., 2016: fig. 6E); Felisacus nigricornis Poppius, 1912 (New Guinea) sclerotized part of ductus seminis curved at Felisacus ochraceus Usinger, 1946 (Guam) least basally and/or medially (e.g., fig. 10H, Felisacus okinawanus Miyamoto, 1965, junior I)...... 3 synonym of F. longiceps 3. Anterior part of C-shaped marking on corium Felisacus ovalau, sp. nov. (Fiji) distinctly inclined posteriorly; labium reach- Felisacus philippinensis Hsiao, 1944 (Philippines) ing posterior margin of metasternum...... Felisacus ponaponensis (Carvalho, 1956) (Micro- ...... F. yasunagai, sp. nov. (fig. 7) nesia) – Anterior part of C-shaped marking on corium Felisacus pulchellus Poppius, 1915, junior syn- directed outward only or slightly inclined onym of F. magnificus posteriorly (fig. 5; F. lindbergae, F. longi- Felisacus rubricuneus Carvalho, 1956 (Micronesia) ceps); labium slightly surpassing posterior Felisacus schuhi, sp. nov. (East Timor, Vanuatu, margin of mesosternum or shorter...... 4 New Caledonia, Solomon Is.) 4. Labium slightly surpassing posterior margin of Felisacus senaru, sp. nov. (Lombok Is.) prosternum, not reaching middle of meso- Felisacus signis Hsiao, 1944, stat. nov. (Borneo) sternum; inner angle of right paramere with Felisacus solomonicus, sp. nov. (Solomon Is.) straight outgrowth (fig. 12M); medial part of Felisacus tanna, sp. nov. (Vanuatu) left paramere without outgrowth (fig. 12N) Felisacus usingeri Woodward, 1954 (Philippines) ...... F. lindbergae, sp. nov. (fig. 5) Felisacus vitilevu, sp. nov. (Fiji) – Labium reaching or almost reaching posterior Felisacus wangae, sp. nov. (South Asia) margin of mesosternum; inner angle of left Felisacus webbi, sp. nov. (Madagascar) paramere with curved outgrowth (figs. 11R, Felisacus yasunagai, sp. nov. (Philippines) 12Q, W, 13AD); medial part of left paramere Felisacus zuparkoi, sp. nov. (Borneo, Philippines) with outgrowth (figs. 11S, 12R, X, 13AE)..5 5. Lateral margins of postocular regions of vertex convex....F. ceylonicus, sp. nov. (fig. 4) – Lateral margins of postocular regions of vertex Key to Felisacus species straight...... 6 1. Corium with distinct, C-shaped, brown to 6. Right paramere almost straight (fig. 12W); api- dark brown marking, reaching R+M anteri- cal part of left paramere widened, flat, with- orly and posteriorly (e.g., fig. 5,F. lindbergae, out outgrowth on dorsal surface (fig. F. longiceps)...... 2 12X)...... F. magnificus Distant (fig. 6) – Marking on corium not reaching R+M anteri- – Right paramere distinctly curved (figs. 12Q, orly, sometimes only in shape of narrow 13AD); apical part of left paramere narrow, marking along inner margin of corium (e.g., outgrowth on dorsal surface of its apical part fig. 5,F. elegantulus, F. jacobsoni, F. mada- present (figs. 12R, 13AE)...... 7 gascariensis)...... 8 7. Outgrowth on inner angle of right paramere 2. Distance between longitudinal sulcus on ver- distinctly curved (fig. 13AD); outgrowth on tex and anterior margin of pronotum shorter middle part of left paramere wide and ser- than eye diameter; vertex distinctly upraised rate (fig. 13AE); vesica with single claw- 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
shaped spicule (fig. 10P)...... 13. Marking on corium reaching R+M posteri- ...... F. zuparkoi, sp. nov. (fig. 7) orly; labium reaching middle of mesoster- – Outgrowth on inner angle of right paramere num; ductus seminis hooked apically (fig. only slightly curved (fig. 12Q); outgrowth 10I)...... F. luzonus, sp. nov. (fig. 5) on middle part of left paramere narrow and – Marking on corium short, not extending smooth (fig. 12R); vesica without spicules toward R+M, labium slightly surpassing (fig. 10H)...... F. longiceps Poppius (fig. 5) posterior margin of mesosternum; ductus 8. Collar not delimited with depression, anterior seminis not hooked apically (fig. 10K)...... part of pronotum distinctly upraised, and ...... F. philippinensis, sp. nov. (fig. 6) covering posterior part of head (Carvalho, 14. Head, pronotum and scutellum red...... 1956: fig. 14)...... F. ponaponensis (fig. 6) ...... F. nigricornis Poppius – Collar more or less delimited, not covering – At least posterior part of pronotum yellow to posterior part of head (Namyatova et al., dark brown...... 15 2016: fig. 4E)...... 9 15. Dorsal side of labial segment II as long as ven- 9. Posterior part of marking on corium subequal tral side (Namyatova et al., 2016: fig. 6E).....16 or wider than eye diameter...... – Dorsal side of labial segment II longer than ven- ...... F. curvatus (fig. 4) tral side (Namyatova et al., 2016:fig. 6D)...... 17 – Marking on hemelytron either short, not extend- 16. Antennal segment I almost straight, only ing towards R+M or its posterior part nar- slightly swollen; vertex upraised; pronotum rower than eye diameter...... 10 with three dark markings posteriorly (fig. 7); 10. Anterior part of marking on corium wide, corium impunctate; inner margin of cuneus subequal to or wider than eye diameter (fig. curved (as in Namyatova et al., 2016: fig. 4, F. cristobalus, F. fedori)...... 11 13E) ...... F. wangae, sp. nov. (fig. 7) – Marking on hemelytron either short, not extend- – Antennal segment I distinctly widened (as in ing towards R+M or its anterior part very Namyatova et al., 2016: fig. 8B); vertex flat narrow, stripelike, distinctly narrower than (as in Namyatova et al., 2016: fig. 6E); pos- eye diameter (e.g., fig. 5, F. filicicola, F. lamb- terior part of pronotum mostly brown, with kinae, F. luzonus)...... 12 yellow to pale brown longitudinal stripe 11. Body length in males 4.4–4.5 mm, in females medially; corium distinctly punctate; inner 4.9–5.9 mm; pronotum yellow brown, more margin of cuneus straight (as in Namyatova or less darkened along posterior margin; et al., 2016: fig. 13F)...... cuneus red (fig. 4); outgrowth on right para- ...... F. crassicornis Usinger (fig. 4) mere rounded (fig. 11AB)...... 17. Posterior part of pronotum brown to dark ...... F. fedori, sp. nov. (fig. 4) brown laterally, with yellow, pale brown or – Body length in male 3.5 mm, in female 4.2 reddish brown longitudinal stripe medially mm; pronotum mostly brown to dark (fig. 5, F. gressitti, F. madagascariensis; fig. 6, brown; with pale brown stripe medially; F. minutus, F. rubricuneus, fig. 7, F. cuneus colorless, with brown margins (fig. vitilevu)...... 18 4); outgrowth on inner part of right param- – Posterior part of pronotum whitish yellow to ere abrupt (fig. 11T)...... pale brown, sometimes with reddish tinge, ...... F. cristobalus, sp. nov. (fig. 4) sometimes with humeral angles or markings 12. Apical half of ductus seminis sclerotized (fig. near humeral angles or stripe along poste- 10I, K)...... 13 rior margin brown to dark brown....22 – Ductus seminis mostly membranous, only 18. Transverse depression on vertex, delimiting small area around ductus seminis sclerotized occipital region present dorsally and later- (e.g., figs. 9, 10N)...... 14 ally (as in Namyatova et al., 2016: fig. 6E); 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 31
cuneus mostly red (fig. 6, F. minutus, F. posteriorly (fig. 11O); vesica as in figure rubricuneus)...... 19 10B...... F. bradi, sp. nov. (fig. 4) – Transverse depression on vertex, delimiting – Pronotum yellow with a pair of brown to dark occipital region present only dorsally; brown markings (fig. 6,F. ovalau; fig. 7,F. cuneus mostly colorless, sometimes with webbi); antennal segment I yellow with red- yellow tinge...... 20 dish tinge or markings; left paramere with 19. Body length in male 2.6; antennal segment II three rounded outgrowths (fig. 13F) or with- red; scutellum yellow; inner part of clavus out outgrowth(s) (fig. 13AA)...... 25 yellow with reddish tinge; medial part of 25. Ventral wall of genital capsule not curved api- right paramere distinctly wider than basal cally (fig. 15N); left paramere narrow and part (fig. 12AC); vesical spicules as in figure sickle shaped, without outgrowths (fig. 9D...... F. minutus Carvalho (fig. 6) 13AA); vesica with long curved spicule, with- – Body length in male 3.4; antennal segment II out toothlike spinules (fig. 10N)...... whitish yellow to yellow, with reddish or ...... F. webbi, sp. nov. (fig. 7) brownish tinge apically; scutellum pale – Ventral wall of genital capsule curved apically brown with reddish tinge; inner part of cla- (fig. 15F); left paramere wide with three vus brown to black; medial part of right rounded outgrowths (fig. 13F); vesica with a paramere as wide as basal part (Carvalho, number of small toothlike spinules only (fig. 1956: fig. 17a); vesical spicules as in figure 9G)...... F. ovalau, sp. nov. (fig. 6) 9H...... F. rubricuneus Carvalho (fig. 6) 26. Distance from eye to pronotum slightly 20. Labium reaching middle of mesosternum; ante- shorter than eye diameter; antennal segment rior part of pronotum brown...... I dark brown with yellow base...... F. madagascariensis Poppius (fig. 5) ...... F. auritulus Distant (fig. 4) – Labium reaching at least posterior margin of – Distance from eye to pronotum slightly longer metasternum or surpassing it; anterior part than eye diameter; antennal segment I yel- of pronotum yellow or pale brown, some- low, brown apically ...... times with reddish tinge...... 21 ...... F. usingeri Woodward (fig. 7) 21. Clypeus mostly dark brown, antennal seg- 27. Anterior part of pronotum brown to dark ment II in male and female ca. 1.6× as long brown...... 28 as head width...... – Pronotum mainly yellow to pale brown, some- ...... F. gressitti Miyamoto (fig. 5) times with brown markings on collar and – Clypeus mostly yellow to pale brown, some- posterior part of pronotum ...... 30 times red; antennal segment II in male ca. 28. Body length in male 4.4–4.7, in female 4.4– 1.9–2.1× and in female ca. 1.9–2.0× as long 4.8; posterior part of pronotum brown to as head width; vesica as in figure 9N...... dark brown; spicules as in figure 9E...... F. vitilevu, sp. nov. (fig. 7) ...... F. nigrescens Carvalho (fig. 6) 22. Depression delimiting occipital region pres- – Body length in males 2.6–3.2 mm, in females ent only dorsally...... 23 2.9–3.5 mm; posterior part of pronotum – Depression delimiting occipital region present mostly yellow to pale brown with brown dorsally and laterally (as in Namyatova et al., markings...... 29 2016: fig. 4E)...... 27 29. Labium reaching posterior margin of meta- 23. Labium reaching abdominal venter...... 24 sternum; antennal segment I yellow to pale – Labium reaching middle of mesosternum....26 brown, sometimes with reddish tinge; scent 24. Pronotum yellow, without brown to dark gland evaporative area uniformly whitish brown markings (fig. 4); antennal segment I yellow; inner part of clavus pale brown to brown; left paramere with single outgrowth brown; cuneus mostly red; middle part of 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
right paramere subequal in length to basal part (fig. 13T); vesical spicules as in figure and apical parts combined (fig. 12I); vesical 9M...... F. tanna, sp. nov. (fig. 7) spicules as in figure 8O...... 33. Collar whitish yellow to yellow with lateral ...... F. lambkinae, sp. nov. (fig. 5) sides often brown to dark brown; left – Labium reaching posterior margin of meso- paramere widened, its apical part twice as sternum or slightly surpassing it; antennal long as wide (fig. 11Q); vesical spicules as segment I reddish with yellow base; scent in figure 8F...F. caledonicus, sp. nov. (fig. 4) gland evaporative area pale brown to – Collar uniformly colored, from whitish yellow brown with red apex; inner part of clavus to brown, often with anterior margin dark- colorless or yellow with red or pale brown ened; left paramere narrow, its apical part ca. margins; cuneus colorless, with brown, 3–5× as long as wide (e.g., figs. 11I, K, 12A orange or red marking along inner margin F)...... 34 and brown margins; middle part of right 34. Darker, pronotum yellow to pale brown with paramere shorter than basal and apical brown transversal stripe posteriorly or at parts combined (fig. 12O); vesical spicules least humeral angles brown; inner part of as in figure 8Q...... F. liui, sp. nov. (fig. 5) clavus pale brown to brown; cuneus at least 30.Labium reaching posterior margin of partly red (fig. 4, F. bau, F. bellus; fig. 6, F. metasternum...... 31 malayensis, F. senaru, F. signis)...... 35 – Labium reaching at most middle of – Paler, usually mostly yellow with clavus and metasternum...... 33 cuneus yellow or colorless; in some species 31. Cuneus mostly red; head and pronotum yel- humeral angles of pronotum brown, clavus low, brown marking on dorsal surface of pale brown and cuneus with red marking, head and along posterior margin of prono- but those characters never present in combi- tum absent; inner part of clavus brown (fig. nation (e.g., fig. 4, F. albus, F australicus, F. 5); scent gland evaporative area brown; vesi- caledonicus; fig. 5, F. filicicola; F. linae, fig. 7, cal spicules as in figure 8M...... F. solomonius, F. tanna)...... 37 ...... F. jacobsoni Poppius (fig. 5) – Cuneus colorless or with yellow tinge; head 35. Medial part of right paramere twice as and pronotum whitish yellow to yellow, wide as basal part (fig. 11J); vesical spic- sometimes with reddish tinge, without ules as in figure 8E...... brown markings; inner part of clavus whit- ...... F. bellus Liu (fig. 4) ish yellow to yellow, sometimes brown – Medial part of right paramere only slightly basally; scent gland evaporative area whitish wider than basal part (figs. 11H, 12Y)...... 36 yellow, sometimes yellow or red apically or 36. Vesical spicules as in figure 8D...... with greenish tinge...... 32 ...... F. bau, sp. nov. (fig. 4) 32. Cuneus transparent with pale brown mar- – Vesical spicules as in figure 9A...... gins; scent gland evaporative area whitish ...... F. malayensis, sp. nov. (fig. 6) yellow, sometimes with greenish tinge; 37. Brown marking along posterior part of pro- medial part of right paramere only slightly notum present; vesical spicules as in figure wider than basal part (fig. 12S); vesical spic- 9I...... F. senaru, sp. nov. (fig. 6) ules as in figure 8R...... – Brown marking along posterior part on prono- ...... F. lordhowensis, sp. nov. (fig. 5) tum absent, sometimes only marking near – Cuneus transparent or with reddish tinge; humeral angle present...... 38 scent gland evaporative area whitish basally 38. Head yellow to pale brown with brown mark- and yellow or red apically; medial part of ing on head dorsally; vesical spicules as in right paramere distinctly wider, than basal figure 9C...... F. myersi, sp. nov. (fig. 6) 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 33
– Head whitish yellow to pale brown, without – Head and pronotum mostly yellow...... 48 marking on head dorsally, sometimes with 47. Cuneus ca. 2.5× as long as wide, mostly red; reddish tinge...... 39 coloration of cuneus varying from transpar- 39. Cuneus at least with yellow tinge or yellow ent with red margins to almost entirely red; apically; sometimes whitish yellow and vesical spicules as in figure 8N...... opaque, sometimes mostly yellow or red...40 ...... F. javanus Reuter (fig. 5) – Cuneus colorless and transparent, sometimes – Cuneus ca. 3× as long as wide, often yellow, brown apically...... 53 sometimes mostly colorless with yellow mark- 40. Medial part of right paramere twice as wide ing along inner margin or yellow with red as basal part (figs. 11A, AF, 12A, C)...... 41 marking along inner margin; vesical spicules – Medial part of right paramere only slightly as in figure 9F...... F. ochraceus Usinger (fig. 6) wider or as wide as basal part (e.g., figs. 11C, 48.Apical part of right paramere swollen (Carv- E, 12K)...... 44 alho, 1956: fig. 17c)...... 41. Cuneus opaque and white, brown apically; ...... F. adamsi Carvalho (fig. 4 ) vesical spicules as in figure 8K...... – Apical part of right paramere not swollen, ...... F. indicus, sp. nov. (fig. 5) evenly tapering toward apex (e.g., figs. 11X, – Cuneus opaque or translucent, at least with yel- Z, 12K, AA, 13R)...... 49 low tinge or yellow apically or along inner 49. Apical part of right paramere distinctly con- margin, sometimes mostly red or yellow...... 42 cave; vesical spicules as in figure 9L...... 42. Labium reaching posterior margin of meso- ...... F. solomonicus, sp. nov. (fig. 7) sternum; vesical spicules as in figure 8A...... – Apical part of right paramere straight or only ...... F. albus, sp. nov. (fig. 4) slightly concave (figs. 11C, X, AD, 12K)...50 – Labium reaching middle of mesosternum....43 50. Labium reaching middle of mesosternum; 43. Cuneus mostly with faint yellow tinge; vesical cuneus entirely red or yellow; vesical spicules spicules as in figure 8J...... as in figure 8B...F. amboinae Woodward (fig. 4) ...... F. glabratus (Motschulsky) (fig. 5) – Labium almost reaching or reaching posterior – Cuneus often yellow at least apically, rarely only with reddish or yellow tinge, spicules as margin of mesosternum; coloration of in figure 8L...... F. insularis Miyamoto (fig. 5) cuneus varying from colorless to uniformly 44. Clavus brown and cuneus red; vesical spic- red or yellow...... 51 ules as in figure 9K.....F. signis Hsiao (fig. 6) 51. Vesica with seven spicules (fig. 8I)...... – Clavus mostly colorless or yellow, sometimes ...... F. filicicola (Kirkaldy) (fig. 5) reddish; rarely brown or pale brown, but in – Vesica with six spicules (figs. 8G, P)...... 52 this case cuneus mostly colorless and trans- 52. Inner part of clavus pale brown; vesical spicules parent or yellow...... 45 as in figure 8P...... F. linae, sp. nov. (fig. 5) 45. Outer margin of medial part of right param- – Inner part of clavus mostly whitish yellow to ere distinctly concave, medial part of right yellow with margin reddish or pale brown; paramere longer than apical part (figs. 11E, vesical spicules as in figure 8G...... 12G, 13C)...... 46 ...... F. dauloi Woodward part 1 (fig. 4) – Outer margin of medial part of right paramere 53. Labium reaching middle of mesosternum or straight or only slightly concave, medial part slightly surpassing it; inner part of clavus of right paramere as long as or shorter than pale brown...... 54 apical part (e.g., fig. 11X, Z, AD)...... 47 – Labium reaching posterior margin of mesoster- 46. Head and pronotum pale brown; vesical spic- num; inner part of clavus often colorless of ules as in figure 8C...... whitish, sometimes pale brown, in this case ...... F. australicus, sp. nov. (fig. 4) humeral angles brown to dark brown...... 55 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
54. Pair of brown to dark brown markings near low. Hemelytron: Mostly colorless and translucent; humeral angles present, distinct; vesical inner part of clavus mostly opaque, with reddish spicules as in figure 9B...... margins; marking along inner margin of corium ...... F. meilingae, sp. nov. (fig. 6) faint, whitish yellow; embolium and cuneus with – Pair of markings near humeral angles absent or reddish margins, embolium yellow apically; mem- faint, mostly pale brown; vesical spicules as brane with whitish-yellow cell. Legs: Yellow with in figure 9J...... F. schuhi, sp. nov. (fig. 6) reddish markings and pale pretarsus. Abdomen: 55. Setae on femora mostly longer than antennal Mostly yellow, genital segments with reddish segment II diameter; humeral angles of pro- marking. SURFACE AND VESTITURE: Dorsum notum often brown to dark brown; antennal and femora clothed with simple setae shorter than segment I often pale brown to brown, rarely antennal segment II diameter; antennal segment I yellow; vesical spicules as in figure clothed with suberect setae subequal to or shorter 8H...... F. elegantulus Reuter (fig. 5) than antennal segment II diameter; abdomen – Setae on femora shorter than antennal segment clothed with suberect setae of different length. II diameter; humeral angles not darkened; STRUCTURE AND MEASUREMENTS: Body antennal segment I often yellow; sometimes 3.9–4.0× as long as pronotum width. Head: pale brown or reddish brown; spicules as in Depression, delimiting occipital region present figure 8G...F. dauloi Woodward part 2 (fig. 4) dorsally and laterally (as in Namyatova et al., 2016: fig. 4E); distance between depression and Felisacus adamsi Carvalho, 1956 pronotum distinctly shorter than eye diameter; longitudinal sulcus on dorsal surface of head lon- Figures 4, 17 ger than eye diameter; distance from eye to pro- Felisacus adamsi Carvalho, 1956: 26 (original notum as long as eye diameter, slightly swollen at description). sides (as in Namyatova et al., 2016: fig. 4E); vertex Diagnosis: Recognized by the following 2.2× as wide as eye; upraised (as in Namyatova et combination of characters: dorsum mostly whit- al., 2016: fig. 6D). Labium (as in Namyatova et al., ish yellow to yellow with reddish markings (fig. 2016: figs. 6D, 9C): Segments I and II strongly 4); antennal segment I cylindrical (as in Namya- reduced, combined less than half as long as seg- tova et al., 2016: fig. 8A), dorsal surface of labial ment III; segment I shorter than wide, segment II segment II elongate posteriorly (Namyatova et slightly longer than wide, its dorsal surface elon- al., 2016: fig. 6D); apical part of right paramere gate posteriorly; segment III slightly shorter than widened; middle part of right paramere less than head length ventrally. Antenna: Segment I cylin- twice as wide as basal part (Carvalho, 1956: fig. drical (as in Namyatova et al., 2016: fig. 8A), 1.6– 17c). 1.8× as long as head width, ca. 0.9× as long as Description: Male. See Carvalho (1956) for pronotum width; segent II ca. 1.6–1.8× as long as the original description. Female. Total length width of head, ca. 1.0–1.1× as long as pronotum 2.9–3.1. COLORATION (fig. 4): Head: Whitish width; segment IV ca. 0.25× as long as segment yellow to yellow, with reddish tinge and markings. III. Thorax: Anterior part of pronotum distinctly Eye brown to dark brown, often with reddish mar- shorter than posterior part; collar delimited; pos- gins. Labium: Yellow. Antenna: Segment I mostly terior part slightly upraised posterior margin of red with yellow base, segments II reddish brown, pronotum straight; pronotum 1.2× as long and ca. segments III–IV brown to dark brown. Pronotum. 1.6× as wide as head; mesoscutum slightly Yellow with pale brown anterior margin, anterior exposed. Hemelytron: Area along inner margin of part with longitudinal red stripe laterally; mesos- corium not swollen; inner margin of cuneus con- cutum yellow; scutellum yellow. Thoracic pleura. vex (as in Namyatova et al., 2016: fig. 13E), medial Yellow; scent gland evaporative area whitish yel- margin of cuneus ca. 3× as long as base. Genitalia 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 35
(from Carvalho, 1956): Genital capsule and Description: Male. Total length 3.7–4.3. Col- aedeagus not examined. Right paramere (Car oration (fig. 4): Head: Mainly whitish yellow to valho, 1956: fig. 17c). Apical part distinct, broad, yellow with reddish tinge and markings, dorsal straight posteriorly; medial part less than twice as surface of head darker anteriorly and along trans- wide as basal part, bearing setae, with outer mar- verse depression. Eye brown with reddish tinge. gin slightly concave and inner margin with dis- Labium: Segments I–III whitish yellow to yellow, tinct swelling; outer angle distinct, widened; inner segment IV yellow. Antenna: Segments I–II yellow angle rounded, without setae; basal part twice with reddish stripes, segments III–IV yellow to shorter than rest of paramere. Left paramere (Car- pale brown, with reddish tinge. Pronotum. Ante- valho, 1956: fig. 17d) paramere L-shaped; apical rior part yellow with whitish yellow collar; ante- part not flattened, with toothlike outgrowth on rior margin and punctures between anterior and posterior side medially and without outgrowth on posterior parts pale brown; stripe laterally reddish; dorsal surface; middle part widened, without posterior part whitish yellow, with brown poste- swelling or outgrowth(s). rior angles; scutellum and mesoscutum whitish Distribution: Micronesia, Ponape Is. (fig. 17). yellow with pale brown punctures between them. Host plants: Unknown. Thoracic pleura. Yellow markings near forecoxa Discussion: F. adamsi has similar external whitish yellow; scent gland evaporative area whit- morphology to several other species, including F. ish yellow with yellow apex. Hemelytron: Mostly auritulus, F. dauloi, F. filicicola, F. insularis, F. translucent, colorless; inner part sometimes with ochraceus, F. schuhi, and F. tanna. (figs. 4–7). We yellow tinge, translucent or opaque, with yellow to have not been able to examine the male genitalia pale brown margins; corium with marking along of this species, because only paratype females inner margin of corium pale brown, darker ante- were available for study from the NMNH. Accord- riorly, not extending toward R+M; embolium ing to the illustrations given by Carvalho (1956), whitish yellow, opaque, with reddish apex and F. adamsi has a unique right paramere with a pale brown outer margin; cuneus whitish yellow broad apical part (Carvalho, 1956: fig. 17c). to yellow with reddish tinge, opaque with pale Material examined: Paratypes: MICRONE- brown apex and brownish margins; membrane SIA: Pohnpei: Ponape Island, Tolotom, 6.83602°N cell whitish yellow. Legs: Coxae whitish yellow; 158.16668°E, 518 m, Jun 1950–Sep 1950, P.A. femora whitish yellow, yellow apically, with red- Adams, 2♀ (00338824; 00341130) (USNM). dish stripe; tibiae whitish yellow to yellow with reddish stripe; tarsi pale brown. Abdomen: Whit- ish yellow, red dorsally. SURFACE AND VESTI- Felisacus albus, sp. nov. TURE: Corium smooth, with shallow and scarce Figures 4, 8A, 11A, B, 14A, 17 punctures. Dorsum with suberect setae longer Diagnosis: Recognized by the following than antennal segment II diameter; antennal seg- combination of characters: pale yellow to white ment I and femora with suberect setae shorter coloration of dorsum with cuneus mostly red than antennal segment II diameter; abdomen (fig. 4), antennal segment I cylindrical (as in clothed with suberect mostly short simple setae. Namyatova et al., 2016: fig. 8A), transverse STRUCTURE AND MEASUREMENTS: Body ca. depression delimiting occipital region extending 4.2–4.3× as long as pronotum width. Head: laterally, vertex upraised, dorsal surface of labial Depression delimiting occipital region present segment II elongate posteriorly (as in Namyatova dorsally and laterally (as in Namyatova et al., et al., 2016: fig. 6D); and unique combination of 2016: fig. 4E); distance between depression and seven vesical spicules, having spicules B and F pronotum distinctly shorter than eye diameter; and lacking spicule A, and having large weekly longitudinal sulcus on dorsal surface longer than sclerotized serrate spicule (fig. 8A). eye diameter; distance from eye to pronotum 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 slightly longer than eye diameter, not swollen lat- outgrowth(s); setae present only on middle part erally (as in Namyatova et al., 2016: fig. 4E); vertex near outer margin. Aedeagus (general view as in ca. 1.5× as wide as eye; upraised (as in Namyatova Namyatova et al., 2016: fig. 22I) conjunctiva et al., 2016: fig. 6D); buccula ca. 0.2–0.25× as long weakly sclerotized; secondary gonopore placed at as clypeus. Labium (as in Namyatova et al., 2016: base of vesica in repose; sclerotization of ductus figs. 6D, 9C): Reaching posterior margin of meso- seminis around secondary gonopore shorter than sternum or slightly surpassing it; segments I and wide; vesica with seven spicules, including spic- II strongly reduced, together less than half as long ules A and F (fig. 8A). as segment II; segment I as long as wide, segment Female. Total length 3.8. COLORATION II slightly longer than wide, its dorsal surface (fig. 4): Similar to male, but dorsal surface of elongate posteriorly; segment III slightly longer head with reddish tinge anteriorly and along than ventral side of head; segment IV ca. 1.5× as transverse depression; posterior part of prono- long as segment III. Antenna: Segment I cylindri- tum whitish yellow with brown posterior mar- cal (as in Namyatova et al., 2016: fig. 8A), ca. 1.4× gin, clavus whitish and transparent with pale as long as head, ca. 1.1–1.2× as long as pronotum brown margins. SURFACE AND VESTITURE: width; segment III slightly longer than segment II; As in male. STRUCTURE AND MEASURE- segment IV ca. 0.3× as long as segment III. Tho- MENTS: Structure as in male; body ca. 3.7× as rax: Anterior and posterior parts subequal in long as pronotum width; vertex in males ca. length; collar distinct; posterior part slightly 1.8× as wide as eye; antennal segment I ca. 1.2× upraised; posterior margin concave; pronotum ca. as long as head width, ca. 0.7× as long as prono- 1.2–1.4× as wide as long and ca. 1.5–1.7× as wide tum width; segment II ca. 1.7× as long as head as head; mesoscutum exposed. Hemelytron: Area width, ca. 1.0× as long as pronotum width; pro- along inner margin of clavus almost flat; inner notum ca. 1.4× as wide as long and ca. 1.8× as margin of cuneus convex (as in Namyatova et al., wide as head. 2016: fig. 13E), outer margin of cuneus ca. 2.5× as Distribution: Known from Taiwan (fig. 17). long as base. Abdomen: Genital capsule rotated Host plants: Collected from Lygodium flex- left at right angle relative to rest of abdomen. Gen- uosum (Schizaeaceae). italia: Genital capsule (fig. 14A) as long as wide, Etymology: The species is named for its very its ventral wall almost twice as long as dorsal wall; pale, almost white coloration, albus in Latin, posterior margin of ventral wall smooth, semi- meaning “white.” oval, without outgrowth, not pointed, not curved Discussion: Female genitalia were not dis- dorsally; its apex inclined rightward; walls of geni- sected. Felisacus albus forms a monophyletic tal capsule not modified; angles of paramere sock- group with F. glabratus, F. indicus and F. insularis ets rounded; distance between paramere sockets (figs. 1–3). All those species are similar to each subequal to half width of genital capsule at base. other in coloration (figs. 4, 5), in the middle part Right paramere (fig. 11A) outer margin of apical of the right paramere more than twice as wide as part slightly concave; medial part more than twice its basal part (figs. 11A, AF, 12A, C), and in the as wide as basal part, bearing setae, with outer shape and configuration of the vesical spicules margin straight and inner margin convex; outer (fig. 8A, J, K, L). Felisacus albus is most similar angle distinct, not widened; inner angle rounded, to F. insularis externally and in morphology of without setae; basal part ca. 0.2–0.25× as long as spicules, including spicules B and F (cf. fig. 8A rest of paramere. Left paramere (fig. 11B) with L). Felisacus insularis differs in having a yel- L-shaped; apical part not flattened, with toothlike low body (fig. 5) and the presence of five vesical outgrowth on posterior side medially (as in fig. spicules (fig. 8K). Felisacus glabratus and F. indi- 11G) and without outgrowth on dorsal surface; cus differ from F. albus in having a mostly yellow middle part widened, without swelling or or colorless cuneus and five vesical spicules. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 37
FIGURE 4. Habitus of Felisacus species. 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 5. Habitus of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 39
FIGURE 6. Habitus of Felisacus species. 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 7. Habitus of Felisacus species.
Material examined: Holotype: THAI- long as base; inner margin middle part of right LAND: Changwat Chiang Mai: Chomtong: Mae paramere less than twice as wide as basal part Klang Waterfall, 18.54944°N 98.56722°E, 28 Nov and its apical part as long as medial part, in outer 1998, A.D. Wright, Lygodium flexuosum (Schiza- margin only slightly concave posteriorly (fig. eaceae), 1♂ (00017861) (AM). Paratypes: THAI- 11C), vesica with four straight spicules, including LAND: Changwat Chiang Mai: Chomtong: Mae spicules A, B, and C (fig. 8B). Klang Waterfall, 18.54944°N 98.56722°E, 28 Nov Redescription: Male. Total length 3.0–3.3. 1998, A.D. Wright, Lygodium flexuosum (Schiza- COLORATION (fig. 4): Head: Mostly pale eaceae), 1♂ (00017860), 1♀ (00017862) (AM). brown, with reddish tinge, ventral side yellow to pale brown. Eye dark brown to black. Labium: Felisacus amboinae Woodward Yellow. Antenna: Segment I yellow with reddish Figures 4, 8B, 11C, D, 14B, 17 tinge, segment II reddish brown; segment III Felisacus amboinae Woodward, 1954: 45 (origi- brown. Thorax: Pronotum and scutellum yellow; nal description). pronotum with pale brown marking close to humeral angle, anterior margin sometimes pale Diagnosis: Recognized by the following brown; thoracic pleura yellow; scent gland combination of characters: mostly yellow color- evaporative area yellow, reddish apically. Hem- ation of dorsum with reddish tinge on cuneus elytron: Mostly translucent, colorless; inner part (fig. 4), cylindrical antennal segment I (as in of clavus with yellow tinge and reddish yellow Namyatova et al., 2016: fig. 8A), transverse margins; marking along inner margin of corium depression on head extending laterally, vertex mostly yellow to pale brown, brown along apical upraised, dorsal surface of labial segment II elon- part of clavus; embolium with pale brown or red gate posteriorly (Namyatova et al., 2016: fig. 6D); margins; cuneus yellow with reddish tinge or body length 3.0–3.3 in male; cuneus ca. 2.5× as margins sometimes reddish brown. Legs: Coxae 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 41 yellow; femora yellow, reddish apically; tibiae 14B) ventral wall ca. 1.7× as long as dorsal wall; mostly reddish yellow, yellow apically; tarsi yel- its posterior margin smooth, semioval, without low with segment III reddish. Abdomen: Mostly outgrowth, rounded, not curved dorsally; apex yellow. SURFACE AND VESTITURE: Body slightly inclined leftward; sides of genital cap- clothed with setae mostly shorter than antennal sule not modified, angles of paramere sockets segment II diameter; dorsum with scarce setae more or less rounded, not projecting; distance mostly shorter than antennal segment II diam- between paramere sockets subequal to half eter; antennal segments I, II and femora clothed width of genital capsule at base; right angle of with suberect setae mostly shorter than width left paramere socket not projecting. Right para- antennal segment II diameter; abdomen clothed mere (fig. 11C) apical part distinct; apex only with short setae. STRUCTURE AND MEA- slightly concave posteriorly; medial part only SUREMENTS: Body ca. 4.1–4.4× as long as pro- slightly wider than basal part, bearing setae, notum width. Head: Depression delimiting with outer margin slightly concave and inner occipital region distinct dorsally and laterally margin convex; outer angle distinct, widened; (as in Namyatova et al., 2016: fig. 4E); distance inner angle rounded, without setae; basal part between depression and pronotum shorter than ca. 0.15–0.2× as long as rest of paramere. Left eye diameter; longitudinal sulcus on dorsal sur- paramere (fig. 11D) L-shaped; apical part not face subequal to eye diameter; distance from eye flattened, with toothlike outgrowth on posterior to pronotum subequal to or slightly longer than side medially and without outgrowth on dorsal eye diameter, not swollen laterally (as in Namy- surface; middle part widened, without swelling atova et al., 2016: fig. 4E); vertex ca. 1.6–1.7× as or outgrowth(s); setae placed only on middle wide as eye, upraised (Namyatova et al., 2016: part near outer margin of paramere. Aedeagus fig. 6D). Labium (as in Namyatova et al., 2016: (general view as in Namyatova et al., 2016: fig. figs. 6D, 9C): Reaching middle of mesosternum; 2IF, G) conjunctiva weakly sclerotized; second- segments I and II strongly reduced; combined ary gonopore placed at base of vesica in repose; less than half as long as segment III; segment II sclerotization of ductus seminis around second- slightly swollen and elongate dorsally; covering ary gonopore shorter than wide; vesica bearing base of segment III; segment III slightly shorter four spicules, including spicules A, B, and C than length of head ventrally; segment IV twice (fig. 8B). as long as segment III. Antenna: Segment I Female. Total length 3.1–3.4. Coloration cylindrical (as in Namyatova et al., 2016: fig. (fig. 4): Similar to male, but antennal segment IV 8A), ca. 1.6–1.7× as long as head width, ca. 1.1– brown; head and pronotum often with reddish 1.2× as long as pronotum width; segment II ca. tinge; tarsal segments I–II yellow and segment 2.0× as long as head width, ca. 1.3–1.4× as long III pale brown to brown or entire tarsi pale as pronotum width. Thorax: Anterior part of brown to brown; scent gland evaporative area as pronotum distinctly shorter than posterior part; in male or whitish yellow, yellow apically, some- collar distinct; posterior part of pronotum only times uniformly yellow; abdomen yellow with slightly upraised; posterior margin of pronotum red markings. SURFACE AND VESTITURE: As slightly concave; pronotum ca. 1.2× as wide as in male. STRUCTURE AND MEASUREMENTS: long and 1.4–1.6× as wide as head. Hemelytron: As in male, antennal segment III subequal to Area along inner margin of corium almost flat; segment II, segment IV ca. 0.7× as long as seg- inner margin of cuneus convex (as in Namya- ment III. Genitalia (general view as in Namya- tova et al., 2016: fig. 13E), outer margin of tova et al., 2016: fig. 23F, G): Dorsal labiate plate cuneus ca. 2.5× as long as base. Abdomen: Geni- wider than distance between apodemes of sec- tal capsule rotated left at right angle relative to ond valvula; mostly smooth, without distinct rest of abdomen. Genitalia: Genital capsule (fig. striations, with semicircular sclerite and distinct 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 8. Vesical spicules of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 43
FIGURE 9. Vesical spicules of Felisacus species. 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 10. A, C–L, O, P. Apical part of aedeagus of Felisacus species B, E, M. Vesica of Felisacus species. N. Aedeagus of F. webbi. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 45
FIGURE 11. Parameres of Felisacus species. 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 12. Parameres of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 47
FIGURE 13. Parameres of Felisacus species. 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 14. Genital capsule of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 49
FIGURE 15. Genital capsule of Felisacus species. 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
FIGURE 16. A, C, E, G, I, K, L, N. Dorsal labiate plate of bursa copulatrix of Felisacus species B, D, F, H, J, M, O. Posterior wall of bursa copulatrix of Felisacus species The small scale is for F. curvatus and F. wangae, the large scale for all others. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 51 sclerotized rings laterally; lateral oviducts placed dark brown; head and pronotum whitish yellow almost medially, very close to each other, sper- to yellow, dorsal surface of labial segment II mathecal gland placed between lateral oviducts; elongate posteriorly (as in Namyatova et al., dorsal labiate plate with distinct tubercles, with- 2016: fig. 6D); head and pronotum mostly yellow out membranous lobe medially. without brown or red markings (fig. 4); body Distribution: Ambon Is., Indonesia (fig. 17). length 4; labium reaching middle of mesoster- Host plants: Unknown. num; distance between eye and pronotum Discussion: Genitalia of the male holotype slightly shorter than eye diameter. and female nontype specimen sampled from the Description (based on a single, partly type locality were examined. Felisacus amboinae destroyed holotype; see also original descrip- is most similar in coloration to F. albus, F. dauloi, tion of Distant, 1912): Total length 4. COLOR- F. filicicola, F. javanus, F. linae, F. ochraceus, and ATION (fig. 4): Head: Yellow to pale brown. F. solomonicus (figs. 4–7), all of them being yel- Eye dark brown to black. Labium: Yellow to low with the cuneus at least partly yellow or red. pale brown. Antenna: Segment I dark brown Among those species, F. dauloi, F. filicicola and with yellow base, segment II dark brown. Tho- F. linae are similar to F. amboinae in shape of the rax: Pronotum and scutellum yellow; thoracic right paramere (cf. fig. 11C with figs. 11D, AD, pleura and scent gland evaporative area yellow. 12K) and distribution, however, those three spe- Hemelytron: Mostly translucent, colorless, cies have an alternative combination of vesical inner part of clavus with yellow tinge. Legs: spicules (cf. fig. 8B with 8G, I, P). Vesical spicules Coxae yellow, femora and tibiae yellow to pale of F. amboinae are most similar to those of the brown, apices of femora and bases of tibiae Timorese species F. linae, with both species hav- somewhat darker, hind tarsus with segment I ing spicules A, B, C, with spicule C distinctly pale brown and segments II–III brown. SUR- posterior in position (cf. fig. 8B with 8P), FACE AND VESTITURE: Setae on dorsum and although the latter species has six spicules. antennal segment I and II mostly shorter than Material examined: Holotype: INDONE- antennal segment II diameter; setae on anten- SIA: Maluku: Ambon Is.: Amboina, 3.66°S nal segments I and II and femora with suberect 128.166°E, no date provided, F. Muir, 1♂ setae mostly shorter than antennal segment II (00400377) (CAS). Paratypes: INDONESIA: diameter. STRUCTURE AND MEASURE- Maluku: Ambon Is.: Amboina, 3.66°S 128.166°E, MENTS: Body ca. 4× as long as pronotum no date provided, F. Muir, 1♂ (00399741), 1♀ width. Head: Depression, delimiting occipital (00399745) (CAS). Additional material: INDO- region present only dorsally; distance between NESIA: Maluku: Ambon Is.: Amboina, 3.66°S depression and pronotum slightly shorter than 128.166°E, no date provided, F. Muir, 4♀ eye diameter; longitudinal sulcus on dorsal (00399742–00399744, 00399746), 2 sex unknown surface shorter than eye diameter; vertex ca. (00399752, 00399756), 1♂ (00399748) (CAS). 3.5× as wide as eye, flat; distance from eye to pronotum slightly shorter than eye diameter, Felisacus auritulus Distant not swollen laterally. Labium (as in in Namya- Figures 4, 17 tova et al., 2016: figs. 6D, 9C): Reaching middle of mesosternum; segments I and II strongly Felisacus auritulus Distant, 1913: 177 (original reduced, combined subequal to half of segment description). III; segment I as long as wide, segment II Diagnosis: Recognized by the following slightly longer than wide, its dorsal elongate combination of characters: antennal segment I posteriorly segment III slightly shorter than distinctly widened (as in Namyatova et al., 2016: length of head ventrally; segment IV twice as fig. 8B), dark brown with yellow base; segment II long as segment III. Antenna: Segment I dis- 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 tinctly swollen (as in Namyatova et al., 2016: Felisacus australicus, sp. nov. fig. 8B), 1.3× as long as head width, 0.6× as long as pronotum width; segment II 2× as long Figures 4, 8C, 11E–G, 14C, 17 as head width, 0.9× as long as pronotum width. Diagnosis: Recognized by the following Thorax: Anterior part of pronotum distinctly combination of characters: relatively dark color- shorter than posterior part; collar distinctly ation with head and pronotum mostly pale delimited; posterior part of pronotum slightly brown without darker markings or stripes; upraised; posterior margin of pronotum slightly antennal segment I pale brown; inner part of concave; pronotum ca. 1.3× as wide as long and clavus pale brown to brown; scent gland evapo- ca. 2.3× as wide as head; mesoscutum slightly rative area pale brown; cuneus matte, yellow or exposed. at least partly red (fig. 4); cylindrical antennal Distribution: Seychelles (fig. 17). segment I (as in Namyatova et al., 2016: fig. Host plants: Unknown. 8A); transverse depression on head extending Discussion: The description of Felisacus auri- laterally, vertex upraised; dorsal surface of labial tulus is based on the holotype, which is seriously segment II elongate posteriorly (as in Namya- damaged, with the abdomen, antennal segments tova et al., 2016: fig. 6D); medial part of right III–IV, most of the hemelytra (except right cla- paramere only slightly wider than basal part vus), and fore- and middle tibiae lost. with outer margin distinctly concave, apical Felisacus auritulus is similar to F. bradi, F. part of right paramere distinctly shorter than ovalau, F. usingeri and F. vitilevu (figs. 4–7) in medial part (fig. 11C); vesica with five spicules, the following characters: broad antennal seg- including spicules A, B, C, H, spicule B with ment I (as in Namyatova et al., 2016: fig. 8B), elongate apical arm (fig. 8C). dorsal surface of labial segment II elongate pos- Description: Male. Total length 3.9. COL- teriorly and pronotum yellow (as in Namyatova ORATION (fig. 4): Head: Yellow to pale brown, et al., 2016: fig. 6D). Felisacus usingeri is most longitudinal sulcus and sides of clypeus brown. similar to F. auritulus, but the latter species dif- Labium: Segments I–III whitish yellow, segment fers in having the distance from eye to prono- IV yellow. Antenna: Segment I pale brown, seg- tum slightly longer than the diameter of an eye, ments II–IV brown to dark brown. Thorax: Pro- antennal segment I is mostly yellow and is notum yellow to pale brown; mesoscutum and known from the Philippines. Felisacus bradi, F. scutellum yellow; thoracic pleura and scent gland ovalau and F. vitilevu differ in having a nar- evaporative area pale brown, slightly paler than rower antennal segment I. Additional charac- pleura. Hemelytron: Inner part of clavus opaque, ters that separate these species from F. auritulus pale brown; outer part of clavus translucent, col- include: F. bradi is known from Tahiti and its orless; corium translucent, colorless with area labium reaches abdominal segment VII; F. ova- along inner margin of corium brown; embolium lau is known from Fiji and antennal segment I opaque, whitish yellow with brown margins; is mostly yellow and the labium reaches abdom- cuneus opaque, whitish yellow with pale brown inal segment V; F. vitilevu is known from Fiji, tinge at outer part and reddish at inner part, mar- and antennal segment I is mostly yellow and the gins brown; membrane translucent with greyish labium reaches the margin of the metasternum tinge; membrane cell pale brown with reddish or slightly surpasses it. tinge. Legs: Yellow to pale brown, tibiae often Material examined: Holotype: SEY- paler apically. Abdomen: Whitish yellow with pale CHELLES: Silhouette, Mare aux Cochons, brown tinge. SURFACE AND VESTITURE: 4.48333°S 55.23333°E, 500 m, Aug 1908, Percy Corium smooth, with shallow punctures. Dor- Sladen Trust Expedition, 1 sex unknown sum with suberect setae mostly longer than (00019490) (BMNH). antennal segment II diameter; antennal segment 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 53
A
F. adamsi F. albus F. amboinae F. auritulus
B
F. australicus F. bau F. bellus F. bismarckensis
C
F. caledonicus F. ceylonicus F. crassicornis F. cristobalus
FIGURE 17. Distribution of Felisacus species. 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
I and femora with suberect setae shorter than sockets subequal to half of genital capsule length antennal segment II diameter; setae on antennal at base; right angle of left paramere socket not segment I scarce; abdomen clothed with suberect, projecting. Right paramere (fig. 11E) apical part mostly short simple setae. STRUCTURE AND distinct, slightly concave posteriorly, twice shorter MEASUREMENTS: Body ca. 4.3× as long as pro- than medial part; medial part slightly wider than notum width. Head: Depression delimiting occip- basal part, bearing setae, its outer margin concave ital region present dorsally and laterally (as in and inner margin convex; outer angle distinct; Namyatova et al., 2016: fig. 4E); distance between inner angle rounded, without setae; basal part ca. depression and pronotum distinctly shorter than 0.15–0.2× as long as rest of paramere. Left para- eye diameter; longitudinal sulcus on dorsal sur- mere (fig. 11F, G) L-shaped; apical part not flat- face longer than eye diameter; distance from eye tened, with toothlike outgrowth on posterior side to pronotum slightly longer than eye diameter, medially (as in fig. 11G) and without outgrowth not swollen laterally (as in Namyatova et al., 2016: on dorsal surface; middle part widened, without fig. 4E); vertex ca. 1.6× as wide as eye, upraised swelling or outgrowth(s); setae placed only on (Namyatova et al., 2016: fig. 6D); buccula ca. middle part near outer margin of paramere. 0.20–0.25× as long as clypeus. Labium (as in Aedeagus conjunctiva weakly sclerotized; second- Namyatova et al., 2016: figs. 6D, 9C): Reaching ary gonopore placed at base of vesica in repose; posterior margin of mesosternum; segments I sclerotization of ductus seminis around second- and II strongly reduced, combined shorter than ary gonopore shorter than wide; vesica with five segment III half; segment I shorter than wide; spicules, including spicules A, B, C, and H, spic- segment II slightly longer than wide, elongate ule B with elongate apical arm (fig. 8C). dorsally; segment III slightly longer than length Female. Total length 4.2–4.4. Coloration of head ventrally; segment IV ca. 1.5× as long as (fig. 4): Head: As in male. Labium: Segments I–III segment III. Antenna: Segment I cylindrical (as in whitish yellow and segment IV pale brown, or Namyatova et al., 2016: fig. 8A), ca. 1.6× as long labium uniformly pale brown. Antenna: Similar to as head width, ca. 1.1× as long as pronotum male, but segment I pale brown with reddish tinge width; segment II ca. 2.0× as long as head width, or brown. Thorax: Similar to male, scent gland ca. 1.3× as long as pronotum width; segments III evaporative area sometimes whitish yellow. Hem- slightly longer than segment II; segment IV ca. elytron: Similar to male, but inner part of clavus 0.3× as long as antennal segment III. Thorax: brown, embolium sometimes pale brown with Anterior part of pronotum only slightly shorter brown margins, cuneus uniformly reddish or than posterior part; collar delimited, posterior whitish. Legs: Similar to male, coxae sometimes part slightly raised; posterior margin concave, whitish yellow. Abdomen: Whitish yellow with pronotum ca. 1.2–1.3× as wide as long and ca. pale brown tinge, sometimes brown ventrally and 1.5–1.6× as wide as head; mesoscutum exposed. dorsally. Surface and vestiture: As in male. STRUC- Hemelytron: Area along inner margin of corium TURE AND MEASUREMENTS: Structure as in not swollen; inner margin of cuneus convex (as males; body ca. 4.1–4.3× as long as pronotum in Namyatova et al., 2016: fig. 13E), outer margin width; vertex ca. 1.3–1.5× as wide as eye; antennal of cuneus ca. 3× as long as base. Genitalia: Geni- segment I ca. 1.4–1.5× as long as head width, ca. tal capsule (fig. 14C) approximately 1.5× as long 0.9–1.0× as long as pronotum width; segment II as wide; ventral wall ca. 1.7× as long as dorsal ca. 1.9–2.2× as long as head width, ca. 1.2–1.5× as wall, posterior margin smooth, without long as pronotum width; pronotum ca. 1.2–1.3× outgrowth(s), slightly tapering, not curved dis- as wide as long and ca. 1.5–1.6× as wide as head. tally; apex inclined leftward; sides of genital cap- Genitalia (as in Namyatova et al., 2016: fig. 23F, sule not modified; angles of paramere sockets G): Dorsal labiate plate wider than distance more or less rounded, distance between paramere between apodemes of second valvula; mostly 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 55 smooth, without distinct striations or longitudinal antennal segment I cylindrical (as in Namyatova membranous ridge, with semicircular sclerites et al., 2016: fig. 8A), depression delimiting and distinct sclerotized rings laterally; lateral ovi- occipital region present dorsally and laterally, ducts placed almost medially, very close to each vertex upraised (as in Namyatova et al., 2016: other, spermathecal gland placed between lateral fig. 6D); body small, length in male 3.2–3.7, in oviducts; dorsal labiate plate with distinct tuber- female 3.8; labium reaching posterior margin of cles, without membranous lobe medially. mesosternum or slightly surpassing it; vesica Distribution: Known from South with six spicules, including spicules A and C, Queensland (Australia) (fig. 17). spicule A shortened (fig. 8D). Host plants: Unknown. Description: Male. Total length 3.5. COLOR- Etymology: The species is named after the ATION (fig. 4): Head: Mostly pale brown, dorsal continent of Australia, where it was collected. surface of head brown anteriorly; longitudinal sul- Discussion: Felisacus australicus is similar in cus brown, maxillary plate whitish yellow, slightly structure to the Micronesian species F. ochraceus paler than surrounding areas. Eye dark brown, and has almost identical male genitalia (cf. figs. reddish at sides. Labium: Uniformly yellow. 11E–G and 13C, D for parameres and figs. 8C Antenna: Segment I brown, yellow basally, seg- and 9F for vesical spicules). However, F. ochra- ment II brown. Thorax: Anterior part of prono- ceus differs from the new species in coloration tum pale brown with brown anterior margin; (fig. 6), possesses a whitish-yellow to yellow head posterior part or pronotum pale brown anteriorly and pronotum, antennal segment I is yellow and brown posteriorly; mesoscutum pale brown, basally and pale brown apically, is sometimes scutellum pale brown, brown apically; thoracic reddish yellow, the inner part of the clavus is pleura brown to dark brown; scent gland evapora- whitish yellow with pale brown or brown mar- tive area dark brown, pale brown basally. Hemely- gins, and the scent gland evaporative area is tron: Mostly translucent, colorless; inner part of whitish yellow. clavus opaque, brown; outer part of clavus brown Material examined: Holotype: AUSTRA- apically; area along inner margin of corium LIA: Queensland: 3 km N on Black Mountain brown; embolium with brown margins and red- Road, 16.66725°S 145.4891°E, 21 Apr 1990, J. dish apex; cuneus red or with brown marking Heraty, 1♂ (00018517) (QM). Paratypes: AUS- along inner margin, margins reddish; anterior TRALIA: Queensland: 3 km N on Black Moun- part of membrane with pale brown shading; tain Road, 16.66725°S 145.4891°E, 21 Apr 1990, membrane cell pale brown, partly reddish. Legs: J. Heraty, 2♀ (00018518, 00018519) (TAMU). Coxae whitish yellow; femora whitish yellow basally and pale brown with reddish tinge apically; Felisacus bau, sp. nov. tibiae pale brown with reddish tinge basally and whitish yellow apically; fore- and middle tarsi yel- Figures 4, 8D, 11H, I, 14D, 17 low to pale brown, hind tarsus whitish yellow. Diagnosis: Recognized by the following Abdomen: Ventral and lateral sides mostly whitish combination of characters: dark coloration, head yellow, genital capsule pale brown with reddish pale brown with brown marking dorsally; pro- tinge, dorsal surface brown. SURFACE AND notum pale brown with dark brown marking VESTITURE: Corium smooth, with shallow and along posterior margin; inner part of clavus scarce punctures. Dorsum, antennal segment I brown; marking along inner margin of corium and femora with suberect setae mostly shorter brown, narrow not extending toward R+M; than antennal segment II diameter; abdomen scent gland evaporative area dark brown, pale clothed with suberect mostly short simple setae. brown basally; cuneus red, sometimes with STRUCTURE AND MEASUREMENTS: Body brown marking along inner margin (fig. 4); 4.2× as long as pronotum width. Head: Depres- 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 sion delimiting occipital region present dorsally cal part not flattened, with toothlike outgrowth on and laterally; distance between depression and posterior side medially (as in fig. 11G) and with- pronotum shorter than eye diameter; longitudinal out outgrowth on dorsal surface; middle part wid- sulcus on dorsal surface longer than eye diameter; ened, without swelling or outgrowth; setae only distance from eye to pronotum slightly longer on middle part near outer margin. Aedeagus (gen- than eye diameter, not swollen laterally (as in eral view as in Namyatova et al., 2016: fig. 22I) Namyatova et al., 2016: fig. 4E); vertex ca. 1.7× as conjunctiva weakly sclerotized; secondary gono- wide as eye, upraised (fig. 6D in Namyatova et al., pore placed at base of vesica; sclerotization of duc- 2016); buccula ca. 0.2–0.25× as long as clypeus tus seminis around secondary gonopore shorter height. Labium (as in Namyatova et al., 2016: figs. than wide; vesica with five spicules, including 6D, 9C): Reaching posterior margin of mesoster- spicules A and C, spicule A shortened, spicule C num or slightly surpassing it; segments I and II large and moved posteriorly (fig. 8D). strongly reduced, combined shorter than half of Female. Total length 3.8. COLORATION, segment III; segment I shorter than wide; segment SURFACE, AND VESTITURE: As in male (fig. II slightly longer than wide, its dorsal surface 4): STRUCTURE AND MEASUREMENTS: elongate posteriorly; segment III slightly longer Structure as in male; body ca. 4.7× as long as than ventral side of head; segment IV 1.5× as long pronotum width; vertex ca. 1.8× as wide as eye; as segment III. Antenna: Segment I cylindrical (as antennal segment I ca. 1.8× as long as head in Namyatova et al., 2016: fig. 8A), ca. 1.7× as long width, ca. 1.2× as long as pronotum width; seg- as head width, ca. 1.1× as long as pronotum width; ment II ca. 2.1× as long as head width, ca. 1.4× segment II ca. 2.1× as long as head width, ca. 1.4× as long as pronotum width; pronotum ca. 1.2× as long as pronotum width. Thorax: Anterior part as wide as long and ca. 1.6× as wide as head. of pronotum only slightly shorter than posterior Distribution: Known from Sarawak, Borneo part; collar delimited; posterior part slightly (Malaysia) (fig. 17). upraised; posterior margin concave; pronotum ca. Host plants: Unknown. 1.2× as wide as long and ca. 1.5× as wide as head; Etymology: The species is named after the mesoscutum exposed. Hemelytron: Area along Bau District, where it was collected. inner margin of corium almost flat; cuneus nar- Discussion: Antennal segments III–IV are lost row, its inner margin convex (as in Namyatova et in the specimens we examined and the anterior al., 2016: fig. 13E), outer margin almost twice as part of the genital capsule is broken. Felisacus bau long as base. Abdomen: Genital capsule rotated is similar to F. bellus, F. jacobsoni, and F. signis in left at right angle relative to the rest of abdomen. appearance (figs. 4 and 6). They all differ from one Genitalia: Genital capsule (fig. 14D) posterior another in the configuration and shape of the vesi- margin smooth, semioval, without outgrowth, not cal spicules (cf. fig. 8D with figs. 8E, M, 9K); in pointed, not curved or inclined, truncate; sides of particular F. bellus possesses an elongate spicule A, genital capsule not modified; margins of paramere and F. jacobsoni and F. signis have only five spic- sockets more or less rounded; distance between ules and lack spicule A. Felisacus jacobsoni can paramere sockets ca. 0.7× as long as genital cap- also be separated by its larger body size, 4.4–4.7 sule width at base. Right paramere (fig. 11H) api- mm in males and 4.6–4.8 mm in females. cal part distinctly curved; apex slightly concave Material examined: Holotype: MALAYSIA: posteriorly; medial part slightly wider than basal Sarawak: Bau District: Bidi, Bau District, part, bearing setae, with outer margin slightly 1.38389°N 110.13357°E, 165 m, 02 Sep 1958, T.C. concave and inner margin convex; outer angle dis- Maa, 1♂ (00043878) (BPBM). Paratype: tinct, not widened; inner angle rounded, without MALAYSIA: Sarawak: Bau District: Bidi, Bau setae; basal part ca. 0.15–0.2× as long as rest of District, 1.38389°N 110.13357°E, 165 m, 02 Sep paramere. Left paramere (fig. 11G) L-shaped; api- 1958, T.C. Maa, 1♀ (00043879) (BPBM). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 57
Felisacus bellus Lin brown, not extending toward R+M anteriorly and continuing to the posterior margin of embo- Figures 4, 8E, 11J, K, 17 lium; embolium reddish apically, margins Felisacus bellus Lin, 2000: 233 (original brown; cuneus red with brown margins, often description). with yellow outer angle; membrane with pale brown or grayish tinge; membrane cell pale Diagnosis: Recognized by the following brown or red. Legs: Coxae whitish yellow; fem- combination of characters: mostly dark color- ora whitish yellow basally and yellow to brown ation, head and pronotum yellow to pale brown, apically; tibiae mostly yellow to pale brown, often with dark brown marking along posterior often coloration gradually changing from brown margin; inner part of clavus brown, sometimes or pale brown basally to yellow or whitish yellow with red outer margin; marking along inner mar- apically; tarsi pale brown, tarsal segment I some- gin of corium brown to reddish brown, not times yellow. Abdomen: Ventral side whitish yel- extending toward R+M anteriorly; cuneus red low, genital capsule pale brown with reddish with brown margins, often with yellow outer posterior margin, lateral and dorsal surfaces angle (fig. 4); antennal segment I cylindrical (as brown. SURFACE AND VESTITURE: Corium in Namyatova et al., 2016: fig. 8A), depression with shallow and scarce punctures. Dorsum delimiting occipital region present dorsally and with suberect setae mostly as long as or slightly laterally; vertex upraised (as in Namyatova et al., longer than antennal segment II diameter; 2016: fig. 6D); body length in male ca. 3.7–4.5, in antennal segment I and femora with suberect female 4.0–4.4; labium reaching posterior mar- setae mostly as long as or slightly shorter than gin of mesosternum; apical part of right para antennal segment II diameter; those on antennal mere distinctly delimited, outer margin of right segment I scarce; abdomen clothed with sub paramere straight (fig. 11J); vesica with six spic- erect mostly short simple setae. STRUCTURE ules, including spicules A, B and G, with spicule AND MEASUREMENTS: Body ca. 4.3–4.4× as A long and narrow (fig. 8E). long as pronotum width. Head: Depression Redescription: Male. Total length 3.7–4.5. delimiting occipital region present dorsally and COLORATION (fig. 4): Head: Mostly yellow to laterally (as in Namyatova et al., 2016: fig. 4E); pale brown, with brown markings and reddish distance from depression to pronotum distinctly tinge; labrum dark brown; labrum dark brown. as long as or longer than eye diameter; longitu- Eye dark brown to black. Labium: Segments I–II dinal sulcus on dorsal surface longer than eye whitish yellow to yellow, segments III–IV yellow diameter; distance from eye to pronotum slightly to pale brown, segment III sometimes with red- longer than eye diameter, not swollen laterally dish tinge. Antenna: Segment I pale brown to (as in Namyatova et al., 2016: fig. 4E); vertex ca. brown, segments II–IV brown to dark brown. 1.7–1.9× as wide as eye; upraised (Namyatova et Thorax: Pronotum yellow with anterior margin al., 2016: fig. 6D); buccula ca. 0.2–0.25× as long and 1/3 of posterior part brown, anterior part as clypeus. Labium (as in Namyatova et al., 2016: sometimes with reddish tinge; mesoscutum pale figs. 6D, 9C): Reaching posterior margin of brown, scutellum pale brown, brown medially mesosternum; segment I and II strongly reduced, and posteriorly; thoracic pleura yellow, meta- combined shorter than half of segment III; seg- pleuron sometimes with brown marking; scent ment I shorter than wide; segment II slightly gland evaporative area whitish yellow basally longer than wide, its dorsal side elongate poste- and brown apically. Hemelytron: Mostly translu- riorly; segment III slightly longer than ventral cent, colorless; inner part of clavus opaque, side of head length; segment IV twice as long as brown, sometimes with red outer margin; area segment III. Antenna: Segment I cylindrical (as along inner margin or corium brown to reddish in Namyatova et al., 2016: fig. 8A), ca. 1.5–1.7× 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 as long as head width, ca. 0.9–1.0× as long as times entire posterior margin brown to dark pronotum width; segment II ca. 2.1× as long as brown or medial marking indistinct; mesoscu- head width, ca. 1.3× as long as pronotum width; tum yellow to pale brown; scutellum yellow to segments III slightly longer than segment II, ca. pale brown, brownish apically, sometimes 0.3× as long as antennal segment IV. Thorax: medial part also brown, rarely with narrow, Anterior part of pronotum slightly shorter than brown longitudinal stripe from center to apex; posterior part; posterior part of pronotum thoracic pleura yellow, sometimes with brown slightly upraised, collar delimited; posterior markings; scent gland evaporative area whitish margin concave, pronotum ca. 1.2–1.3× as wide yellow basally, yellow to brown apically, some- as long and ca. 1.7× as wide as head; mesoscu- times mostly brown. Hemelytron: Similar to tum exposed. Hemelytron: Area along inner male, but inner part of clavus sometimes yellow margin of corium almost flat; inner margin of or pale brown with brown margins; area along cuneus convex, outer margin of corium ca. 2.5– inner margin of corium sometimes short; not 3× as long as base. Abdomen: Genital capsule reaching R+M; cuneus rarely mostly whitish yel- rotated left at right angle relative to the rest of low with reddish tinge with pale brown margins abdomen. Right paramere (fig. 11J) apical part and red apical third. Legs: Similar to male, fem- distinct; apex slightly concave posteriorly; ora sometimes uniformly yellow, middle tibia medial part less than twice as wide as basal part, rarely pale brown, tibiae sometimes with pale bearing setae, with outer margin straight and brown apex. Abdomen: Ventral and lateral sides inner margin swollen; outer angle distinct; inner whitish yellow, lateral side often with pale brown angle rounded, without setae; basal part ca. to brown markings, segment IX pale brown, or 0.15–0.2× as long as rest of paramere. Left para- whitish yellow with reddish apex, dorsal surface mere (fig. 11K) L-shaped; apical part not flat- whitish yellow to brown; rarely abdomen uni- tened, with toothlike outgrowth on posterior formly whitish yellow with reddish apex. SUR- side medially (as in fig. 11G) and without out- FACE AND VESTITURE: As in male. growth on dorsal surface; middle part widened, STRUCTURE AND MEASUREMENTS: Struc- without swelling or outgrowth; setae only on ture as in male; body ca. 3.9–4.4× as long as pro- middle part near outer margin. Aedeagus (as in notum width; vertex ca. 1.9–2.2× as wide as eye; fig. 8E) conjunctiva weakly sclerotized; second- antennal segment I ca. 1.3–1.7× as long as head ary gonopore placed at base of vesica; sclerotiza- width, ca. 0.8–1.0× as long as pronotum width; tion of ductus seminis around secondary segment ca. II 1.8–2.1× as long as head width, gonopore shorter than wide; vesica bearing six ca. 1.1–1.2× as long as pronotum width; prono- spicules, including spicules A, B and G, with tum ca. 1.2–1.3× as wide as long and ca. 1.7× as spicules A long and narrow (fig. 8E). wide as head. Genitalia (as in Namyatova et al., Female. Total length 4.0–4.4. COLORATION 2016: fig. 23F, G): Dorsal labiate plate wider than (fig. 4): Head: Yellow, often anterior and ventral distance between apodemes of second valvula; surfaces, buccula, mandibular, maxillary plates mostly smooth, without distinct striations or and clypeus whitish yellow to yellow; frons, dor- longitudinal membranous ridge, with semicircu- sal and lateral sides and clypeus basally some- lar sclerites and distinct sclerotized rings later- times with reddish tinge. Eye dark brown to ally; lateral oviducts placed almost medially, black. Labium: Similar to male, sometimes uni- very close to each other, spermathecal gland formly yellow. Antennae. As in male. Thorax: placed between lateral oviducts; dorsal labiate Pronotum yellow with pale brown anterior mar- plate with distinct tubercles, without membra- gin, often with reddish tinge laterally, posterior nous lobe medially. margin with markings medially and near Distribution: China, Taiwan, Vietnam humeral angles brown to dark brown, some- (fig. 17). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 59
Host plants: Unknown. Felisacus bismarckensis, sp. nov. Discussion: We examined one female para- type and a digital image of the holotype that is Figures 4, 10A, 11L, M, 14E, 17 deposited in the Taiwan Agriculture Research Diagnosis: Recognized by the following Institute. We were therefore unable to examine combination of characters: dark brown to black the male genitalia. However, the additional spec- C-shaped marking on corium, extending toward imens we were able to examine are very similar R+M anteriorly and posteriorly, its anterior part to the types and also were collected in southeast not inclined posteriorly (fig. 4); head not swollen Asia (Vietnam and China), and we concluded laterally behind eyes; vertex upraised; antennal that they are conspecific with F. bellus. segment I widened (as in Namyatova et al., 2016: Felisacus bellus is almost indistinguishable fig. 8B); right paramere without outgrowth on from F. jacobsoni in coloration (fig. 4), having inner angle (fig. 11L); and at least half of ductus similar size and body structure, but the latter seminis sclerotized, straight apically (fig. 10A). species differs in the labium reaching the poste- Description: Male. Total length 3.1. COL- rior part of the metasternum and differs in the ORATION (fig. 4): Head: Mostly yellow to pale morphology of the vesical spicules (cf. figs. 8E brown, buccula and clypeus brown apically. Eye with 8M); in particular, F. jacobsoni possesses reddish brown. Labium: Segments I–II brown, only five spicules with spicule 1 broad. segment III pale brown, yellow apically, segment Felisacus bellus is also similar to F. bau in IV yellow. Antenna: Segment I reddish brown external morphology, coloration (fig. 4) and in with yellow base; segment II brown, segments the number of spicules (cf. figs. 8D with 8E), but III–IV pale brown. Thorax: Anterior part yellow the latter species is lacking spicules B and G and to pale brown, with anterior margin and marking possesses spicule C. Felisacus bau also has the near forecoxa brown; posterior part brown; right paramere with the apical part not delimited mesoscutum and scutellum uniformly brown, (fig. 11H). scutellum sometimes with reddish apex; thoracic Material examined: Paratype: TAIWAN: pleura and scent gland evaporative area brown. Nantou: Tungpu, Nantou Hsien, 23.91448°N Hemelytron: Clavus opaque, uniformly brown; 120.6812°E, 1200 m, 20 Jun 1983–24 Jun 1983, corium, embolium and cuneus mostly translu- K.C. Chou and C.Y. Wong, 1♀ (00005119) cent, colorless; corium with C-shaped brown (NMNS). Additional material: CHINA: Yun- marking, reaching R+M anteriorly and posteri- nan: Longyang, 35.16666°N 117.25°E, 1600 m, orly, anterior part of this marking not inclined 12 Aug 2006, unknown, 1♀ (00017868) posteriorly; embolium with margins and apex (NKMU). Menghai, 21.97145°N 100.44914°E, brown; cuneus yellow apically, with outer margin 05 Oct 1979, Le-Yi Zheng, 1♀ 00018515) brown and inner margin colorless; membrane (NKMU). THAILAND: Nakhon Ratchasima: translucent, gradually changing color from pale Wang Nam Khieo: Sakaerat Environmental brown anteriorly and whitish yellow posteriorly; Research Station, 14.5075°N 101.9275°E, 410 m, membrane cell pale brown to brown. Legs: Fore- 19 Aug 2008–21 Aug 2008, Yasunaga and coxa brown, middle and hind coxae yellow with Shishido, 1♀ (00021582) (TYCN). VIETNAM: pale brown to brown markings medially; fore- Hòa Bình: Hoa Binh: Near Mai Chau [Mai and middle femora yellow apically, sometimes Chon, Mai Chou], 20.667°N 105.084°E, 04 Nov with reddish or pale brown marking; hind femur 1990, Belokobylskij, 2♂ (00018447), 1♀ yellow with yellow, pale brown or red apex and (00018514) (ZISP). Lam Dong: 6 km S Dalat, pale brown or red marking at apical half; tibiae 11.91724°N 108.44464°E, 1450 m, 09 Jun 1961– yellow, red or pale brown basally; tarsi yellow to 07 Jul 1961, N.R. Spencer, 2♀ (00017873, pale brown. Abdomen: Ventral side yellow, lateral 00271514) (BPBM). side brown basally and reddish apically; dorsal 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 surface brown. SURFACE AND VESTITURE: genital capsule width at base. Right paramere Corium smooth, with shallow, scarce punctation. (fig. 11L) distinctly curved apically; apex slightly Dorsum, antennal segment I and femora clothed concave, medial part narrower than basal part, with setae shorter than antennal segment II without setae, outer margin of medial part con- diameter; abdomen clothed with long erect setae. vex, inner margin concave, not widened; outer STRUCTURE AND MEASUREMENTS: Body angle absent; inner angle present, rounded, with- ca. 4.0× as long as pronotum width. Head: Trans- out outgrowth, bearing setae; basal part slightly verse depression delimiting occipital region pres- shorter than rest of paramere. Left paramere (fig. ent only dorsally, not extending laterally 11M) apical part slightly flattened, with toothlike (Namyatova et al., 2016: fig. 6E), distance outgrowth apically, without outgrowth dorsally; between depression and pronotum shorter than middle part widened dorsally, inner margin of eye diameter; longitudinal sulcus on dorsal sur- middle part swollen, without setae and out- face of head shorter than eye diameter; eye diam- growth. Aedeagus (general view as in Namyatova eter slightly longer than distance between eye et al., 2016: fig. 22M) vesica without spicules; and pronotum; vertex ca. 2.1× as wide as eye; apical half of ductus seminis sclerotized, straight, vertex upraised. Labium: Reaching posterior secondary gonopore placed near phallotheca margin of metasternum; segments I and II mouth in repose (fig. 10A). shorter than width, combined subequal to half of Female. Total length 3.5–3.7. COLORATION segment III; dorsal surface of segment II not (fig. 4):Head: Similar to male, but clypeus some- elongate posteriorly (as in Namyatova et al., times uniformly brown, eye dark brown. Labium: 2016: fig. 6E); segment III as long as ventral side As in male. Antenna: Segment I brown to reddish of head length; segment IV twice as long as seg- brown with yellow base, segment II brown, seg- ment III. Antenna: Segment I as long as head ments III and IV pale brown to brown. Thorax: width, widened (as in Namyatova et al., 2016: fig. Similar to male; metapleuron sometimes pale 8B), ca. 1.0× as long as head width, ca. 0.6× as brown with brown posterior margin. Abdomen: long as pronotum width; segment II ca. 1.7× as Yellow ventrally and brown dorsally and apically. long as head width, ca. 1.1× as long as pronotum STRUCTURE AND MEASUREMENTS: Struc- width; segment III slightly shorter than segment ture as in male; body ca. 4.0–4.1× as long as pro- II and ca. 1.5× as long as segment IV. Thorax: notum width; vertex ca. 2.1–2.2× as wide as eye; Anterior part of pronotum shorter than posterior antennal segment I ca. 1.0–1.1× as long as head part; collar delimited; posterior part of pronotum width, ca. 0.6–0.7× as long as pronotum width; slightly upraised; posterior margin of pronotum segment II ca. 1.7× as long as head width, ca. straight or concave; pronotum ca. 1.3× as wide as 1.0× as long as pronotum width; pronotum ca. long and ca. 1.6× as wide as head; mesoscutum 1.2–1.3× as wide as long and ca. 1.6–1.7× as wide slightly exposed or not exposed. Hemelytron: as head. Genitalia (as in fig. 16A, B): Dorsal labi- Area along inner margin of corium swollen; ate plate very small and transparent, as wide as cuneus with inner margin straight (Namyatova distance between apodemes of second valvulae, et al., 2016: fig. 13F), outer margin of cuneus without striations or membranous ridge medi- twice as long as base. Genitalia: Genital capsule ally; semicircular sclerite and sclerotized rings (fig. 14E) ventral wall ca. 1.3× as long as dorsal absent; lateral oviducts placed in anterior part; wall, with posterior margin of ventral wall oval, spermathecal gland attached near anterior mar- slightly acute with apex placed almost medially, gin; dorsal labiate plate without distinct tuber- not curved dorsally, without toothlike outgrowth; cles, without membranous lobe posteriorly. sides of genital capsule not modified; margins of Distribution: Bismarck Archipelago (New paramere sockets distinctly acute; distance Britain Is.) (fig. 17). between paramere sockets subequal to half of Host plants: Unknown. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 61
Etymology: The species is named after Bis- behind eye, longitudinal stripe pale brown, buc- marck Archipelago, where it was collected. cula, frons pale brown with reddish tinge near Discussion: The genital capsule was not clypeus base, clypeus mostly brown, whitish yel- examined. The external morphology of Felisacus low only basally. Eye dark brown. Labium: Seg- bismarckensis is similar to F. lindbergae, F. longi- ments I–II whitish yellow, segment III yellow, ceps, F. magnificus, and F. zuparkoi (figs. 4–7), segment IV pale brown basally and gradually but those four species differ in having a flat ver- darkened to brown apically. Antenna: Brown, tex (as in Namyatova et al., 2016: fig. 6E), a dis- segment I whitish yellow basally. Thorax: Ante- tinct outgrowth on the inner angle of the right rior part of pronotum whitish yellow, posterior paramere (cf. fig. 11L with figs. 12M, Q, W, part of pronotum yellow; mesoscutum and scu- 13AD) and the ductus seminis is distinctly sickle tellum yellow with reddish tinge; thoracic pleura shaped apically (cf. fig. 10A with 10F, H, J, O). and scent gland evaporative area whitish yellow. Material examined: Holotype: PAPUA Hemelytron: Mostly translucent, whitish yellow; NEW GUINEA: East New Britain Province: clavus with yellow tinge and brown depression Gazelle Peninsula, Gaulim, 4.45°S 142.08333°E, delimiting part of clavus; area along inner mar- 04 Nov 1962, R.W. Strandtmann, 1♂ (00043207) gin of corium brown, narrow and short, not (BPBM). Paratypes: PAPUA NEW GUINEA: extending toward R+M; embolium with outer East New Britain Province: Gazelle Peninsula, margin partly dark brown; membrane with pale Gaulim, 4.45°S 142.08333°E, 04 Nov 1962, R.W. brown tinge; membrane cell pale brown. Legs: Strandtmann, 1♂ (00043208), 2♀ (00043209, Coxae whitish yellow; femora whitish yellow 00043210), sex unknown (00043211) (BPBM). basally and yellow to pale brown apically; tibiae yellow to pale brown; tarsi brown. Abdomen: Whitish yellow. SURFACE AND VESTITURE: Felisacus bradi, sp. nov. Corium smooth, with shallow and dense punc- Figures 4, 10B, 11N, O, 14F, 16C, D, 21 tures. Dorsum, antennal segment I and femora Diagnosis: Recognized by the following clothed with suberect setae shorter than anten- combination of characters: antennal segment I nal segment II diameter; abdomen with suberect slightly widened; head and pronotum whitish mostly short simple setae. STRUCTURE AND yellow to yellow, without brown markings (fig. MEASUREMENTS: Body ca. 4.3× as long as 4); antennal segment I mostly brown; dorsal pronotum width. Head: Depression delimiting surface of labial segment II elongate posteriorly occipital region present only dorsally, not (as in Namyatova et al., 2016: fig. 6D); labium extending laterally (Namyatova et al., 2016: fig. reaching abdominal segment VII; ventral wall 6E); distance between depression and pronotum of genital capsule not curved apically (fig. 14F); as long as eye diameter; longitudinal sulcus on left paramere widened with large outgrowth dorsal surface shorter than eye; distance from posteriorly (fig. 11O); sclerotization of ductus eye to pronotum slightly shorter than eye diam- seminis around secondary gonopore elongate, eter, not swollen dorsally; vertex ca. 1.8× as wide but distinctly shorter than remaining part of as eye; vertex slightly raised; buccula ca. 0.2– ductus seminis, dorsal wall of the sclerotized 0.25× as long as clypeus. Labium: Reaching portion of ductus seminis longer than ventral abdominal segment VII; segments I and II wall; vesica with a number of small tooth- reduced, subequal to half of segment III; seg- shaped sclerites apically and large sclerotized ment I shorter than wide; segment II slightly areas (fig. 10B). longer than wide, its dorsal surface elongate Description: Male. Total length 4.3. COL- posteriorly (as in Namyatova et al., 2016: fig. ORATION (fig. 4): Head: Mostly whitish yellow 6D); segment III slightly longer than length of to yellow, dorsal and lateral sides pale brown head ventrally; segment IV ca. 0.3× as long as 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 segment III. Antenna: Segment I slightly longer vesica with number of small toothlike sclerites than head width, slightly widened medially and two large sclerites (fig. 10B). (Namyatova et al., 2016: fig. 8B), ca. 1.3× as long Female. Total length 4.9. COLORATION (fig. as head width, ca. 0.8× as long as pronotum 4): Head: Mostly yellow to pale brown, ventral width; segment II ca. 2.0× as long as head width, side yellow, apical part of clypeus brown; eyes ca. 1.2× as long as pronotum width; segments III dark brown. Labium: As in male. Antennae. shorter than segment II; segment IV ca. 0.25× as Brown to dark brown, segment I yellow basally. long as antennal segment III. Thorax: Anterior Thorax: Anterior part of pronotum whitish yel- part of pronotum distinctly shorter than poste- low, posterior part of pronotum mostly pale rior part; posterior part slightly upraised; poste- brown; mesoscutum and scutellum as in male; rior margin concave, pronotum ca. 1.3× as wide mesopleuron yellow; metapleuron and scent as long and ca. 1.6× as wide as head; mesoscu- gland evaporative area whitish yellow with tum exposed. Hemelytron: Area along inner greenish tinge. Legs: Similar to male, but tarsal margin of corium almost flat; inner margin of segments I and II of fore- and middle tibia pale corium convex (as in Namyatova et al., 2016: fig. brown. Hemelytron: As in male. Abdomen: Whit- 13E), outer margin of corium ca. 3× as long as ish yellow with green tinge. SURFACE AND base. Abdomen: Genital capsule rotated left at VESTITURE: As in male. STRUCTURE AND right angle relative to rest of abdomen. Genita- MEASUREMENTS: Structure similar to male, lia: Genital capsule (fig. 14F) twice as long as but labium reaching abdominal segment VIII; wide; ventral wall ca. 1.3× as long as dorsal wall, body ca. 4.2× as long as pronotum width; vertex its posterior margin smooth, semioval, without ca. 2.0× as wide as eye; antennal segment I ca. outgrowth, apex slightly inclined to left-hand 1.0× as long as head width, ca. 0.6× as long as side, rounded, not curved dorsally; sides of geni- pronotum width; segment II ca. 1.7× as long as tal capsule not modified; margins of paramere width of head, ca. 1.0× as long as width of pro- sockets angulate; distance between paramere notum; pronotum ca. 1.4× as wide as long and sockets ca. 0.3× as long as width of genital cap- ca. 1.6× as wide as head. Genitalia (fig. 16C, D): sule at base. Right paramere (fig. 11N) curved in Dorsal labiate plate wider than distance between apical part, but apical part only weakly delim- apodemes of second valvula; with striated longi- ited; apex straight posteriorly; medial part only tudinal membranous ridge, with semicircular slightly wider than basal part, its outer margin sclerites and distinct sclerotized rings laterally; straight, inner margin slightly widened, bearing lateral oviducts placed almost medially, shifted setae; outer angle present, very shallow; inner posteriorly, very close to each other, spermathe- angle rounded, not bearing setae or outgrowth; cal gland placed between lateral oviducts; dorsal basal part ca. 0.25–0.3× as long as rest of para- labiate plate with distinct tubercles anteriorly, mere. Left paramere (fig. 11O) L-shaped, wid- without membranous lobe medially. ened; apical part not flattened, with large Distribution: Tahiti (fig. 21). outgrowth on posterior side medially and with- Host plants: Unknown. out outgrowth on dorsal surface; middle part Etymology: This species is named after Brad widened, without swelling or outgrowth(s); Balukjian, who provided material for this study. setae present only posteriorly. Aedeagus con- Discussion: Antennal segment IV is lost in junctiva sclerotized only basally, secondary the male. Felisacus bradi is most similar to F. gonopore placed at base of vesica; sclerotized auritulus, F. usingeri, and F. ovalau in the com- part of ductus seminis around secondary gono- bination of the broad antennal segment I and pore elongate, but distinctly shorter than rest of the dorsal surface of labial segment II is elongate ductus seminis; dorsal wall of sclerotized part of posteriorly (as in Namyatova et al., 2016: fig. ductus seminis longer than its ventral wall; 6D). Felisacus usingeri is known from the Philip- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 63 pines and differs by antennal segment I yellow, pale brown to dark brown, often with reddish brown apically, and the labium reaching the tinge laterally. Labium: Whitish yellow to yellow, middle of the mesosternum. Felisacus ovalau is segment I sometimes somewhat darker than known from Fiji and differs by the antennal seg- other segments, segment III sometimes with ment I yellow with a reddish tinge, the prono- reddish stripe ventrally. Antenna: Segment I yel- tum with pair of brown markings near the low to pale brown often darker apically, some- humeral angles, the left paramere is broad with times with reddish tinge; segment II varying three rounded outgrowths (fig. 13F), and sclero- from almost yellow to pale brown or reddish tization of the ductus seminis at the secondary brown, sometimes with reddish tinge, often gonopore is broad and almost as long as wide darker than segment II; segments III–IV brown (fig. 9G). Felisacus auritulus is known from the to dark brown. Thorax: Pronotum whitish yel- Seychelles and differs by antennal segment I low to yellow with anterior margin pale brown being distinctly broad and the labium reaches to brown, often lateral sides of collar brown; col- the middle of the mesosternum. lar rarely with reddish tinge; posterior part of Material examined: Holotype: FRENCH pronotum sometimes with pair of brown mark- POLYNESIA: Society Islands: Moorea Is.: Uufau ings on posterior margin at distance from Pass, 17.536°S 149.8697°W, 420 m, 15 Mar 2009, humeral angles; mesoscutum whitish to yellow, Peter Oboyski, 1♂ (00399753) (UCB). Paratype: scutellum whitish to whitish yellow, sometimes FRENCH POLYNESIA: Society Islands: Moorea somewhat paler than mesoscutum; thoracic Is.: ridge between Mt Mouaroa and Mt Ruahite, pleura yellow to pale brown, sometimes with near Mt Mouaroa overlook, 17.54794°S pale brown to brown marking; scent gland evap- 149.84955°W, 488 m, 11 Dec 2008, C.P. Ewing, orative area whitish yellow, rarely with greenish 1♀ (00399754) (UCB). tinge. Hemelytron: Mostly translucent, colorless; inner part of clavus sometimes yellow with yel- low to pale brown margins; area along inner Felisacus caledonicus, sp. nov. margin of corium whitish yellow to pale brown; Figures 4, 8F, 11P, Q, 14G, 17 embolium often with yellow apex and yellow to Diagnosis: Recognized by the following pale brown margins; margins of cuneus yellow combination of characters: dorsum mostly to brown; membrane cell yellow. Legs: Coxae whitish yellow to yellow coloration with lateral whitish yellow; femora whitish yellow basally sides of collar often brown; humeral angles and yellow apically, rarely uniformly yellow, whitish yellow (fig. 4), cylindrical antennal seg- sometimes with red apices; tibia uniformly whit- ment I (as in Namyatova et al., 2016: fig. 8A), ish yellow to yellow, sometimes with red bases; transverse depression delimiting occipital tarsi pale brown, often segment I and sometimes region present dorsally and laterally, vertex segment II whitish to yellow. Abdomen: Ventral upraised (as in Namyatova et al., 2016: fig. 6D); and lateral side of abdomen whitish yellow, dor- labium reaching posterior margin of mesoster- sal surface of pregenital segments and entire num; cuneus ca. 3× as long as base; apical part genital capsule yellow to pale brown, sometimes of left paramere widened, twice as long as wide abdomen uniformly yellow to pale brown. SUR- (fig. 11Q); vesica with four spicules, including FACE AND VESTITURE: Corium smooth, spicules A and B (fig. 8F). often with very shallow, indistinct punctures. Description: Male. Total length 3.5–3.8. Dorsum, antennal segment I, femora and abdo- COLORATION (fig. 4): Head: Mostly whitish men clothed with setae mostly subequal to or yellow to yellow, sometimes pale brown dorsally, somewhat shorter than antennal segment II sometimes with reddish tinge and markings; diameter, those setae sometimes; abdomen longitudinal sulcus yellow to pale brown. Eye clothed with erect short setae subequal to or 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 shorter than antennal segment II diameter. socket slightly acute and left one rounded; dis- STRUCTURE AND MEASUREMENTS: Body tance between paramere sockets ca. 0.4× as long ca. 4.3–4.6× as long as pronotum width. Head: width of genital capsule at base. Right paramere Depression, delimiting occipital region present (fig. 11P) apical part distinct, straight; medial dorsally and laterally (as in Namyatova et al., part only slightly less than twice as wide as basal 2016: fig. 4E); distance between depression and part, bearing row of setae, with outer margin pronotum distinctly shorter than eye diameter; straight or slightly concave, and inner margin longitudinal sulcus on dorsal surface of head swollen; outer angle present; inner angle subequal to eye diameter; distance from eye to rounded, without setae or outgrowth; basal part pronotum longer than eye diameter, not swollen ca. 0.5× as long as rest of paramere. Left param- laterally (as in Namyatova et al., 2016: fig. 4E); ere (fig. 11Q) widened, L-shaped, only slightly vertex ca. 1.8–2.1× as wide as eye diameter; curved; apical part straight, twice as long as upraised (Namyatova et al., 2016: fig. 6D). wide, with toothlike outgrowth on posterior side Labium (as in Namyatova et al., 2016: figs. 6D, medially (as in fig. 11G); middle part widened, 9C): Reaching posterior part of mesosternum or without outgrowth; setae only on middle part slightly surpassing it; segments I and II reduced, near outer margin. Aedeagus (general view as in combined shorter than half of segment III; seg- Namyatova et al., 2016: fig. 22I) conjunctiva ment I shorter than wide; segment II slightly weakly sclerotized; secondary gonopore placed longer than wide, its dorsal surface elongate at base of vesica in repose; sclerotization of duc- posteriorly; segment III subequal to length of tus seminis around secondary gonopore shorter head ventral side; segment IV ca. 1.5× longer than wide; vesica with four spicules, including A than segment III. Antenna: Segment I cylindri- and B (fig. 8F). cal (as in Namyatova et al., 2016: fig. 8A), ca. Female. Total length 3.7–4.0. COLORATION 1.5–1.9× as long as head width, ca. 1.0–1.3× as (fig. 4): Head: Similar to male, sometimes head long as pronotum width; segment II ca. 2.1–2.3× uniformly pale brown. Labium: Similar to male, as long as head width, ca. 1.4–1.5× as long as rarely uniformly pale brown. Antenna: Similar to pronotum width; segment III slightly longer male, coloration of segment I varying from yel- than segment II; segment III ca. 0.3× as long as low to pale brown, sometimes darker apically or segment IV. Thorax: Anterior part of pronotum with reddish tinge. Thorax: Similar to male, scent slightly shorter than posterior part; collar delim- gland evaporative area whitish yellow. Legs and ited; posterior part slightly upraised; posterior hemelytron. As in male. Abdomen: Whitish yel- margin slightly concave, pronotum ca. 1.1–1.3× low ventrally and yellow to pale brown dorsally, as wide as long and ca. 1.4–1.6× as wide as head; sometimes segment IX uniformly yellow to pale mesoscutum exposed or not exposed. Hemely- brown, sometimes abdomen uniformly whitish tron: Area along inner margin of corium almost yellow to pale brown. SURFACE AND VESTI- flat; inner margin of cuneus convex (as in TURE: As in male. STRUCTURE AND MEA- Namyatova et al., 2016: fig. 13E), outer margin SUREMENTS: Structure as in male; body ca. of cuneus ca. 3× as long as base. Abdomen: Gen- 4.2–4.4× as long as pronotum width; vertex ca. ital capsule rotated left at almost right angle 2.0–2.3× as wide as eye diameter; antennal seg- relative to rest of abdomen. Genitalia: Genital ment I ca. 1.7–1.8× as long as head width, ca. capsule (fig. 14G) almost as long as wide, ventral 1.0–1.1× as long as pronotum width; segment II wall ca. 1.5× as long as dorsal wall, with poste- ca. 2.1–2.3× as long as head width, ca. 1.2–1.4× rior margin of ventral wall rounded, smooth, as long as pronotum width; pronotum ca. 1.2– not curved, without outgrowth(s); apex of geni- 1.4× as wide as long and ca. 1.6–1.7× wider than tal capsule slightly inclined rightward; sides of head. Genitalia (as in Namyatova et al., 2016: fig. genital capsule not modified; right paramere 23F, G): Dorsal labiate plate wider than distance 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 65 between apodemes of second valvula; mostly N.L.H. Krauss, 1♂ (00042280) (BPBM). Mt. smooth, without distinct striations, with semicir- Koghi, 22.16666°S 166.53333°E, 525 m, Dec cular sclerite and distinct sclerotized rings later- 1983, N.L.H. Krauss, 2♀ (00042287, 00042288) ally; lateral oviducts placed almost medially, very (BPBM). Yahoué, 22.19686°S 166.6655°E, Feb close to each other, spermathecal gland placed 1976, N.L.H. Krauss, 1♀ (00042293), 1♂ between lateral oviducts; dorsal labiate plate with (00042284) (BPBM). Province des Iles Loyauté: distinct tubercles, without membranous lobe Fayaoue, Ouvea Is., 20.65774°S 166.54238°E, 25 medially. m, Dec 1968, N.L.H. Krauss, 1♂ (00042306) Distribution: New Caledonia, Loyalty Is. (BPBM). New Caledonia, 1940, unknown col- (fig. 17). lector, 1♂ (00042304) (BPBM). Host plants: Unknown. Etymology: The species is named after New Felisacus ceylonicus, sp. nov. Caledonia from where the type material was collected. Figures 4, 10C, 11R, S, 14H, 17 Discussion: Felisacus caledonicus is very sim- Diagnosis: Recognized by the following com- ilar to F. elegantulus in external morphology (fig. bination of characters: marking on corium dark 4) but the latter species can be separated by the brown, C-shaped, its anterior and posterior parts following combination of characters: the lateral reaching R+M (fig. 4); cuneus brown apically; margins of the collar are yellow; the humeral antennal segment I widened (as in Namyatova angles of the pronotum are often brown to dark and Cassis, in press: fig. 8B), red; antennal seg- brown; the apical part of the left paramere is not ment II yellow with red markings; vertex swollen broad, ca. 3× as long as wide; and the presence laterally behind eye; ventral wall of genital capsule of five vesical spicules, including spicule B. with angulate outgrowth posteriorly (fig. 14H); Material examined: Holotype: NEW right paramere sickle shaped, with inner angle CALEDONIA: Province Sud: Yahoué, bearing distinctly curved outgrowth; inner margin 22.19686°S 166.6655°E, 20 Feb 1963, C.M. of medial part of right paramere convex, without Yoshimoto, 1♂ (00042285) (BPBM). Paratypes: swelling (fig. 11R); apical part of left paramere NEW CALEDONIA: Province Nord: Hiengh- narrow, with tooth apically and outgrowth dor- ene, 20.68793°S 164.9434°E, 80 m, 14 Aug sally; medial part with narrow outgrowth (fig. 1979–17 Aug 1979, G.M. Nishida, 2♂ 11S); distal half of ductus seminis sclerotized (fig. (00042282, 00042283), 4♀ (00042289– 10C); vesica without spicules (fig. 10C). 00042292) (BPBM). Province Sud: Col de la Description: Male. Total length 2.6. COL- Pirogue, 22.08161°S 166.3165°E, 330 m, 14 Feb ORATION (fig. 4): Head: Mostly pale brown with 1963, N.L.H. Krauss, 4♀ (00042298–00042300, reddish markings around eyes and antennal fos- 00042303) (BPBM). Col de la Pirogue, sae and on clypeus. Eye red. Labium: Segments 22.08161°S 166.3165°E, 23 Jan 1962, N.L.H. I–II pale brown; segments III–IV yellow. Antenna: Krauss, 1♂ (00042286) (BPBM); 14 Feb 1963, Segment I red; segment II yellow with red mark- N.L.H. Krauss, 4♀ (00042296, 00042297, ings; segments III–IV pale brown. Thorax: Ante- 00042301, 00042302) (BPBM). Crouen, rior part of pronotum brown; posterior part of 21.53333°S 165.88333°E, Mar 1959, N.L.H. pronotum dark brown; scutellum and mesoscu- Krauss, 2♀ (00042294, 00042295) (BPBM). Les tum dark brown; thoracic pleura brown, scent Bois de Sud, 22.17°S 166.76°E, 200 m, 23 Apr gland evaporative area whitish yellow. Hemely- 2005, Cassis, Wall, Tatarnic, Monteith, 1♂ tron: Clavus opaque, dark brown; corium, embo- (00039957) (AM). Monts des Koghis, 22.15151°S lium, and cuneus mostly translucent and 166.53394°E, 500 m, Jan 1969, N.L.H. Krauss, colorless; corium with brown anterior angles and 2♂ (00042279, 00042281) (BPBM); Feb 1973, C-shaped dark brown marking with anterior and 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 posterior parts of this marking reaching costal along inner margin of corium swollen; inner margin; embolium and cuneus brown apically; margin of cuneus straight (as in Namyatova et al., membrane and membrane cell brown. Legs: 2016: fig. 13F), outer margin of cuneus twice as Coxae whitish yellow; femora mostly whitish yel- long as base. Abdomen: Genital capsule rotated low, fore- and middle femora reddish apically, left 45° relative to the rest of abdomen. Genitalia: hind femur with apical half reddish brown; tibiae Genital capsule (fig. 14H). Twice as long as wide; mostly brown with apical 1/3 whitish yellow; tarsi ventral wall ca. 1.5× longer than dorsal wall, with whitish yellow. Abdomen: Yellow, segments VIII– apex not curved dorsally; with angulate out- IX brown. SURFACE AND VESTITURE: Corium growth at left side and with right side folded; left smooth with distinct punctation. Dorsum with side of genital capsule not widened; margins of setae mostly longer than antennal segment II paramere sockets angulate; distance between diameter; antennal segment I clothed with sub paramere sockets ca. 0.3× as long as genital cap- erect setae shorter than antennal segment II sule width at base. Right paramere (fig. 11R) diameter; femora with suberect setae twice as sickle shaped, medial part narrower than basal long as antennal segment II diameter; abdomen part, without setae, only slightly curved, its outer with erect setae of different length. STRUCTURE margin straight; inner margin concave without AND MEASUREMENTS: Body ca. 3.9× as long swelling; outer angle of right paramere absent, as pronotum width. Head: Depression, delimiting inner angle present, with curved outgrowth and occipital region, present only dorsally; distance bunch of setae on inner margin, basal part ca. between depression and pronotum slightly longer 0.5× as long as rest of paramere. Left paramere than eye diameter; longitudinal sulcus on dorsal (fig. 11S) curved medially; apical part not flat- surface distinctly shorter than eye diameter; dis- tened, with toothlike outgrowth apically and with tance from eye to pronotum distinctly longer outgrowth on dorsal surface; inner margin of than eye diameter, swollen laterally; vertex ca. middle part with narrow outgrowth, bearing 1.6× as wide as eye diameter, flat (Namyatova et setae. Aedeagus vesica without spicules; half of al., 2016: fig. 6E); buccula ca. 0.2× as long as clyp- ductus seminis sclerotized, secondary gonopore eus. Labium: Almost reaching posterior margin placed near phallotheca mouth in repose; ductus of mesosternum; segments I and II reduced, com- seminis hooked apically (fig. 10C). bined slightly longer than half of segment III; Female. Total length 3.5. COLORATION (fig. dorsal surface of segment II not elongate posteri- 4): Head: Similar to male, but maxillary plate orly (as in Namyatova et al., 2016: fig. 6E); seg- with reddish marking and eyes yellow with red- ment III shorter than ventral side of head; dish tinge. Labium: Uniformly yellow. Antenna: segment IV twice as long as segment III. Antenna: As in male. Thorax: Similar to male, thoracic Segment I widened medially (as in Namyatova et pleura pale brown, mesopleuron with brown al., 2016: fig. 8B), ca. 1.3× as long as head width, marking. Hemelytron: As in male. Legs: Hind ca. 0.7× as long as pronotum width; segment II femur mostly reddish brown with basal 1/3 whit- ca. 1.8× as long as head width, ca. 0.9× as long as ish yellow. SURFACE AND VESTITURE: As in pronotum width; antennal segment III slightly male. STRUCTURE AND MEASUREMENTS: longer than segment II and twice as long as seg- Structure as in male; body ca. 3.9× as long as ment IV. Thorax: Anterior part of pronotum dis- pronotum width; vertex ca. 2.2× as wide as eye tinctly shorter than posterior part; collar distinct; diameter; segment I 1.4× as long as head width, posterior part of pronotum distinctly upraised; 0.7× as long as pronotum width; segment II 1.7× posterior margin of pronotum usually straight, as long as head width, 0.8× as long as pronotum sometimes slightly concave; pronotum ca. 1.1– width; pronotum ca. 1.2× as wide as long and ca. 1.2× as wide as long and ca. 1.9× as wide as head; 2.1× as wide as head. Female genitalia were not mesoscutum slightly exposed. Hemelytron: Area investigated. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 67
Distribution: Sri Lanka (fig. 17). slightly surpassing middle of mesosternum; Host plants: Unknown. occipital region present only dorsally; distinctly Etymology: The species is named after the protruding eye; hemelytron with distinct punc- former name of Sri Lanka, “Ceylon,” where it was tures; right and left parameres sickle shaped collected. (Carvalho, 1956: fig. 17h, I). Discussion: Felisacus ceylonicus is similar to F. Redescription: Male. No specimens were lindbergae, F. longiceps, F. magnificus,and F. zupar- examined. See Usinger (1946) and Carvalho koi (figs. 4–7) in the pattern of coloration and the (1956) for description of male. structure of the ductus seminis (fig. 10C, F, H, J, P). Female. Total length 3.3–3.5. COLORATION Felisacus longiceps also has similar paramere struc- (fig. 4): Head: Whitish yellow to pale brown, buc- ture (cf. fig. 11R, S with fig. 12Q, R). All aforemen- cula anteriorly, clypeus apically and tubercle tioned species differ from F. ceylonicus in having around antennal fossa often pale brown. Eye the vertex straight laterally behind the eye. Felisacus brown, sometimes with reddish tinge. Labium: magnificus can also be separated from F. ceylonicus Uniformly yellow. Antenna: Segment I yellow by the right paramere straight and the left paramere dark brown apically and often pale to brown flattened.Felisacus zuparkoi also differs from F. cey- basally; segment II yellow basally and pale brown lonicus in having claw-shaped vesical spicules, and apically; segments III–IV brown to dark brown, F. zuparkoi differs in the labium not surpassing the rarely segment III basally or entire segment IV posterior margin of the prosternum. pale brown. Thorax: Anterior part of pronotum Material examined: Holotype: SRI LANKA: yellow to pale brown, often with anterior margin Central Prov.: Peradeniya, Univ. Peradeniya, pale brown; punctures between anterior and pos- 7.25416°N 80.60194°E, 335 m, 20 Mar 1999, TJ terior parts pale brown; posterior part of prono- Henry, A Wijesekara and C Ariyadasa, 1♂ tum pale brown to brown with yellow or pale (00399955) (USNM). Paratype: SRI LANKA: brown stripe medially; scutellum and mesoscu- Central Prov.: Peradeniya, Univ. Peradeniya, tum whitish yellow to yellow with punctures 7.25416°N 80.60194°E, 335 m, 20 Mar 1999, TJ between them pale brown; thoracic pleura whit- Henry, A Wijesekara and C Ariyadasa, 1♀ ish yellow to yellow; scent gland evaporative area (00399956) (USNM). whitish yellow to yellow. Hemelytron: Clavus opaque, its inner part yellow to brown, outer part Felisacus crassicornis Usinger, 1946 yellow to pale brown; corium whitish, translu- cent, areas around punctures opaque; area along Figures 4, 17 inner margin of corium pale brown to brown Felisacus crassicornis Usinger, 1946: 70 (original narrow and short, not extending toward R+M; description). embolium whitish yellow to yellow, translucent Diagnosis: Recognized by the following to opaque, with yellow margins; cuneus translu- combination of characters: pronotum mostly cent, colorless with outer margin yellow; mem- brown with yellow medial longitudinal stripe; brane translucent, colorless with pale brown or marking along inner margin of corium pale grayish tinge; membrane cell pale brown. Legs: brown to brown, short and narrow, not extend- Yellow, bases of coxae and femora often whitish ing toward R+M (fig. 4), antennal segment I yellow; tibia sometimes pale brown apically, tar- mostly yellow, dark brown apically; antennal sal segments I and II often pale brown. Abdomen: segment I widened (as in Namyatova and Cas- Yellow with pale brown dorsal surface. SUR- sis, in press: fig. 8B); dorsal surface of labial seg- FACE AND VESTITURE: Corium often with ment II not elongate posteriorly (as in distinct punctures. Dorsum and femora anteri- Namyatova et al., 2016: fig. 6E); labial segment orly clothed with suberect setae mostly longer III shorter than ventral side of head; labium than antennal segment II diameter; setae on dor- 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 sum scarce; antennal segment I with rare sub and females in his redescription of the species. erect setae shorter than antennal segment II We had only females available for this study. diameter; abdomen clothed with suberect mostly Felisacus crassicornis can be confused with F. short simple setae. STRUCTURE AND MEA- auritulus externally (fig. 4); the latter species SUREMENTS: Body ca. 3.6–3.9× as long as pro- differs by the antennal segment I dark brown notum width. Head: Depression delimiting with a yellow base, the dorsal surface of labial occipital region present only dorsally; distance segment II elongate posteriorly, covering anten- between depression and pronotum distinctly nal segment II, and the punctures on the hem- longer than eye diameter; longitudinal sulcus on elytra absent or indistinct. The parameres of F. dorsal surface shorter than eye diameter; dis- crassicornis are most similar to those of F. tance from eye to pronotum slightly longer than ponaponensis (cf. Carvalho, 1956: fig. 17h with eye diameter, not swollen; vertex ca. 2.1–2.5× as fig. 13I, J). These two species are also similar in wide as eye diameter; upraised; buccula ca. 0.2– having a pale body, without any dark stripes or 0.25× as long as clypeus. Labium: Reaching pos- markings on the hemelytra. However, F. ponapo- terior margins of mesosternum but not nensis differs in the collar not being delimited surpassing it; segments I and II reduced, com- posteriorly, and the pronotum covering the bined slightly longer than half of antennal seg- head to the depression delimiting the occipital ment III; segment I shorter than wide; segment region. We have no information on the struc- II slightly longer than wide, subequal to half of ture of the aedeagus of F. crassicornis. segment III, its dorsal surface not elongate pos- Material examined: Holotype: GUAM: teriorly (as in Namyatova et al., 2016: fig. 8B); Machanao, 25 Nov 1936, O.H. Swezey, 1♀ segment III shorter than length of ventral side of (BPBM_TCN 00002954) (BPBM). Paratypes: head; segment IV subequal to half of segment GUAM: Machanao, 05 Jun 1936, R.L. Usinger, III. Antenna: Segment I only slightly longer than 1♀ (BPBM_TCN 00002955) (BPBM). Additional head width, widened (as in Namyatova et al., material: GUAM: Mariana Is.: Guam, Mt. 2016: fig. 8B), ca. 0.9–1.1× as long as head width, Lamlam, Dec 1958, N.L.H. Krauss, 1♀ ca. 0.6–0.7× as long as pronotum width; segment (00043260) (BPBM). Upi Trail, 05 May 1936, II ca. 1.2–1.5× as long as head width, ca. 0.8– O.H. Swezey, 1♀ (00043252) (BPBM). Yigo, 1.0× as long as pronotum width; segment III dis- 13.53194°N 144.88027°E, 126 m, Oct 1957, tinctly longer than segment II; segment IV ca. N.L.H. Krauss, 6♀ (00043253–00043258) 0.4–0.5× as long as segment III. Thorax: Anterior (BPBM); Feb 1958, N.L.H. Krauss, 1♀ part of pronotum shorter than posterior part; (00043259) (BPBM). collar distinct; posterior part of pronotum slightly upraised; posterior margin of pronotum Felisacus cristobalus, sp. nov. straight or concave; pronotum ca. 1.3–1.4× as wide as long and ca. 1.5–1.6× as wide as head; Figures 4, 10D, 11T, U, 14I, 17 mesoscutum usually exposed, sometimes not Diagnosis: Recognized by the following exposed. Hemelytron: Area along inner margin combination of characters: marking on hemely- of corium swollen; inner margin of cuneus tron dark brown, with anterior part widened and straight, outer margin of cuneus twice as long as not reaching R+M (fig. 4); antennal segment I base. Female genitalia were not examined. widened (as in Namyatova et al., 2016: fig. 8B), Distribution: Mariana Is. (fig. 17). mostly brown to dark brown; cuneus mostly Host plants: Ferns (Usinger, 1946). translucent, colorless; posterior part of pronotum Discussion: Usinger (1946) described F. mostly brown to dark brown with pale brown crassicornis from two females, housed in the stripe medially; inner angle of right paramere BPBM. Carvalho (1956) also examined males with short and abrupt outgrowth (fig. 11T); out- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 69 growth on medial part of left paramere widened, STRUCTURE AND MEASUREMENTS: Body angulate (fig. 11U); half of ductus seminis sclero- ca. 4.1× as long as pronotum width. Head: Trans- tized, straight, sickle shaped apically (fig. 10D). verse depression delimiting occipital region pres- Description: Male. Total length 3.5. COL- ent only dorsally, not extending laterally ORATION (fig. 4): Head: Mostly yellow to pale (Namyatova et al., 2016: fig. 6E), distance brown, frontal side slightly paler, buccula anteri- between depression and pronotum slightly orly and clypeus apically brown. Eye brown with shorter than eye diameter; longitudinal sulcus on reddish tinge. Labium: Segment I yellow to pale dorsal surface of head shorter than eye diameter; brown; segment II brown; segment III yellow, distance from head to pronotum distinctly lon- pale brown ventrally; segment IV yellow. ger than eye diameter; vertex ca. 1.6× as wide as Antenna: Segment I pale brown basally and eye, flat (Namyatova et al., 2016: fig. 6E).Labium: brown to dark brown apically, segment II brown Reaching or slightly surpassing posterior margin to dark brown; segments III–IV pale brown to of prosternum; segments I and II combined as brown, sometimes somewhat paler than segment long as segment III; segment I twice as long as II. Thorax: Anterior part of pronotum pale brown wide; segment II slightly longer than wide, with dark brown anterior margin; posterior part slightly shorter than segment I, its dorsal surface of pronotum dark brown with yellow or pale not elongate posteriorly (as in Namyatova et al., brown longitudinal sulcus medially; mesoscu- 2016: fig. 6E); segment III distinctly shorter than tum and scutellum pale brown to brown; tho- ventral side of head length; segment IV ca. 1.5× racic pleura pale brown, with brown to dark as long as segment III. Antenna: Segment I brown markings; scent gland evaporative area slightly longer than head width, swollen basally whitish yellow. Hemelytron: Mostly translucent, (as in Namyatova et al., 2016: fig. 8B), ca. 1.2× as only clavus opaque; inner part of clavus brown long as head width, ca. 0.8× as long as pronotum to dark brown; outer part of clavus pale brown to width; segment II ca. 1.8× as long as head width, brown; corium translucent, colorless, with mark- ca. 1.2× as long as pronotum width; segment III ing along inner margin of corium dark brown, its slightly longer than segment II; segment IV sub- anterior part widened, wider than eye diameter, equal to half of segment III. Thorax: Anterior not reaching R+M vein, posterior part of mark- part of pronotum shorter than posterior part; ing not widened, reaching costal vein; embolium collar distinct; posterior part slightly upraised; colorless with brown margins; cuneus colorless posterior margin of pronotum slightly concave; with brown outer margin; membrane brown pronotum ca. 1.1× as wide as long and ca. 1.5× anteriorly and colorless posteriorly; membrane as wide as head; mesoscutum usually exposed. cell brown. Legs: Coxae whitish yellow; femora Hemelytron: Area along inner margin of corium whitish yellow, often yellow apically, with pale swollen; inner margin of cuneus straight (as in brown or brown marking in apical half and often Namyatova et al., 2016: fig. 13F), outer margin of pale brown apically; foretibia pale brown with cuneus twice as long as base. Genitalia: Genital whitish yellow apex; middle and hind tibiae pale capsule (fig. 14I) ventral wall ca. 1.5× as long as brown basally and whitish yellow apically; tarsal dorsal wall, with posterior margin rounded, not segments I whitish yellow; segments II–III yel- curved dorsally, with toothlike outgrowth; mar- low to pale brown. SURFACE AND VESTI- gin of paramere sockets distinctly angulate; dis- TURE: Corium smooth, with rare and shallow tance between paramere sockets ca. 0.25× as long punctation. Dorsum and forefemur with setae as genital capsule width at base. Right paramere slightly longer than antennal segment II diame- (fig. 11T) distinctly curved; in apical half; apex ter; antennal segment I, middle and hind femora slightly concave posteriorly; medial part slightly clothed with suberect setae shorter than antennal wider than basal part, without setae, outer mar- segment II; abdomen with short erect setae. gin of medial part convex, inner margin concave 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 without swelling; outer angle absent; inner angle Discussion: Antennal segment IV is lost in present with abrupt swelling, bearing setae; basal the female. Felisacus cristobalus is similar to F. part of paramere almost as long as rest of para- fedori in the marking on the corium, with the mere. Left paramere (fig. 11U) apical part not anterior part not reaching R+M and is broad (fig. flattened, with toothlike outgrowth on posterior 4): The latter species differs in having a yellow side apically and with outgrowth on dorsal sur- pronotum, brown to dark brown posteriorly, the face; middle part with wide outgrowth, bearing cuneus is red, and the outgrowth on the right setae. Aedeagus vesica without spicules; half of paramere is acute (fig. 11AB). ductus seminis sclerotized, secondary gonopore Material examined: Holotype: SOLOMON placed near phallotheca mouth; ductus seminis ISLANDS: San Cristobal: Kira-Kira, 10.44138°S hooked apically (fig. 10D). 161.91666°E, 25 m, 05 Nov 1964, R. Straatman, Female. Total length 4.2. COLORATION 1♂ (00043871) (BPBM). Paratypes: SOLOMON (fig. 4): Head: As in male. Labium: Similar to ISLANDS: Savo Is., West Coast, 9.13333°S male, but labial segment I pale brown. Antenna: 159.78333°E, 50 m, 19 Oct 1970, N.L.H. Krauss, Similar to male, but segment I brown, same 1♀ (00043873) (BPBM). San Cristobal: Kira color as segment II. Thorax: Similar to male, but Kira, 10.44138°S 161.91666°E, 100 m, Jan 1976, posterior part of pronotum dark brown with yel- collector unknown, 1♀ (00043872) (BPBM). low stripe medially, mesoscutum brown; scutel- lum pale brown. Hemelytron and legs: As in male. Abdomen: Ventral whitish lateral sides, Felisacus curvatus Hu and Zheng, 2001 segment IX and dorsal surface brown. SUR- Figures 4, 10E, 11V, W, 14J, 16L, M, 18 FACE AND VESTITURE: Similar to male, abdomen clothed with erect setae mostly as long Felisacus curvatus Hu and Zheng, 2001: 417 as or shorter than antennal segment I width. (original description). STRUCTURE AND MEASUREMENTS: Struc- ture as in male; body ca. 4.3× as long as prono- Diagnosis: Recognized by the following com- tum width; vertex ca. 1.7–1.9× as wide as eye; bination of characters: marking along inner mar- antennal segment I ca. 1.1–1.3× as long as head gin of corium brown, with anterior part not width, ca. 0.8× as long as pronotum width; seg- reaching R+M not widened, and posterior part ment II ca. 1.7–1.8× as long as head width, ca. reaching R+M and noticeably widened (fig. 4); 1.1–1.2× as long as pronotum width; pronotum depression delimiting occipital region distinct ca. 1.2× as wide as long and ca. 1.5–1.6× as wide only dorsally; segment I cylindrical (as in Namya- as head. Genitalia (as in fig. 16A, F): Dorsal labi- tova et al., 2016: fig. 8A); head and pronotum yel- ate plate very small and transparent, as wide as low to pale brown; right paramere sickle shaped, distance with apodemes of second valvulae, with outer angle rounded and inner angle distinct, without striations or membranous ridge medi- without outgrowth (fig. 11V); sclerotization of ally; semicircular sclerite and sclerotized rings ductus seminis around secondary gonopore short, absent; lateral oviducts placed in anterior part; twice as long as wide; vesica with three serrate spermathecal gland attached near anterior mar- spicules, bearing toothlike outgrowths (fig. 10E). gin; dorsal labiate plate without distinct tuber- Description: Male. Total length 4.2–4.8. cles, without membranous lobe posteriorly. COLORATION (fig. 4): Head: Mainly yellow to Distribution: Solomon Is. (fig. 17). pale brown, with reddish tinge; buccula, tubercle Host plants: Unknown. around eye, marking below antennal fossa, man- Etymology: The species is named after San dibular and maxillary plates whitish yellow; clyp- Cristobal Is., where two specimens out of three eus brown apically. Eye dark brown with reddish were collected. tinge. Labium: Whitish yellow to yellow. Antenna: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 71
Segment I whitish yellow to yellow, reddish api- erect setae. STRUCTURE AND MEASURE- cally, sometimes uniformly whitish yellow; seg- MENTS: Body 4.2–4.6× as long as pronotum ment II yellow basally and pale brown to brown width. Head: Depression, delimiting occipital apically, sometimes uniformly pale brown to region, present only dorsally; distance between brown; segments II–IV pale brown to brown. Tho- depression and pronotum as long as eye diameter; rax: Anterior part of pronotum yellow to pale longitudinal sulcus on vertex shorter than eye brown with anterior margin brown, often with diameter; distance from eye to pronotum dis- reddish tinge; posterior part of pronotum often tinctly longer than eye diameter, not swollen; ver- yellow to pale brown often with humeral angle tex ca. 1.7–2.2× as wide as eye, flat (as in fig. 7E). brown, rarely posterior part of pronotum uni- Labium: Slightly surpassing posterior margin of formly dark brown; mesoscutum yellow to pale prosternum; segment I twice as long as than wide, brown, rarely with reddish tinge, sometimes with segment II slightly longer than wide, its dorsal brown marking medially, rarely uniformly brown surface not elongate posteriorly (as in Namyatova to dark brown; scutellum yellow to pale brown, et al., 2016: fig. 6E); segments I and II combined rarely with reddish tinge; rarely brown to dark almost subequal to segment III; segment III brown; thoracic pleura pale brown to dark brown; shorter than ventral side of head, segment IV ca. scent gland evaporative area whitish yellow basally 1.5× as long as segment III. Antenna: Segment I and brown apically. Hemelytron: Mostly translu- cylindrical (as in Namyatova et al., 2016: fig. 8A), cent and colorless, clavus opaque; inner part of ca. 1.4–1.5× as long as head width, ca. 0.8× as long clavus pale brown to brown, sometimes with red- as pronotum width; segment II ca. 1.7–2.0× as dish tinge; outer part of clavus, brown, often long as head width, ca. 0.9–1.1× as long as prono- darker than inner part; marking along inner mar- tum width; segments III distinctly longer than gin of corium reddish brown or brown; its ante- segment II; segment IV subequal to half of seg- rior part very short not reaching R+M and ment III. Thorax: Anterior part of pronotum posterior part reaching R+M, twice as wide as shorter than posterior part; collar delimited; pos- middle part; embolium brown basally, often yel- terior part slightly upraised; posterior margin of low to pale brown apically with margins partly pronotum straight or concave; pronotum ca. 1.0– pale brown to brown; cuneus reddish or reddish 1.2× as wide as long and ca. 1.8–2.0× as wide as brown apically; membrane pale brown to brown; head; mesoscutum usually exposed. Hemelytron: membrane cell brown, often with reddish tinge. Area along inner margin of corium swollen; inner Legs: Coxae whitish yellow, femora whitish yellow margin of cuneus convex (as in Namyatova et al., basally and yellow apically, sometimes with red- 2016: fig. 13E), outer margin of cuneus ca. 3× as dish tinge apically; hind and middle tibiae some- long as base. Abdomen: Genital capsule rotated times with reddish-brown marking on apical half; left at small angle relative to rest of abdomen. tibiae whitish yellow to yellow, sometimes with Genitalia: Genital capsule (fig. 14J) twice as long reddish tinge basally; tarsi whitish yellow to yel- as wide; ventral wall ca. 1.3× as long as dorsal low. Abdomen: Whitish yellow to yellow ventrally wall, its posterior margin smooth, without out- and laterally, genital capsule pale brown, striped growths; semioval, slightly pointed, curved dor- laterally and dorsal surface reddish, rarely abdo- sally from right side; left side of genital capsule men uniformly reddish. SURFACE AND VESTI- not widened; margins of socket slightly acute; dis- TURE: Corium smooth, with rare and shallow tance between paramere sockets subequal to half punctation. Dorsum and femora with setae dis- of genital capsule width at base. Right paramere tinctly longer than antennal segment II diameter; (fig. 11V) distinctly curved in apical half; apex antennal segment I clothed with suberect setae straight posteriorly; medial part slightly wider subequal to or slightly longer than antennal seg- than basal part, without setae, outer margin of ment II diameter; abdomen clothed with short medial part convex; inner margin convex; outer 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
A
F. curvatus F. dauloi F. elegantulus F. fedori
B
F. filicicola F. glabratus F. gressitti F. indicus
C
F. insularis F. jacobsoni F. lambkinae F. linae
FIGURE 18. Distribution of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 73 angle rounded; inner angle distinct, without out- near posterior margin, posterior to lateral ovi- growths and setae; basal part of right parameres. ducts; dorsal labiate plate with membranous 0.7× as long as rest of paramere. Left paramere semicircular lobe anteriorly, with tubercles only (fig. 11W) L-shaped; apical part not flattened, on this lobe and around it. without toothlike outgrowth apically or medially Distribution: China, Laos, Vietnam (fig. 18). and without outgrowth on dorsal surface; middle Host plants: Unknown. part slightly widened, without swelling or out- Discussion: Felisacus curvatus was described growth; setae placed on middle part near outer by Hu and Zheng (2001) from China (Hainan). margin. Aedeagus (fig. 10E) conjunctiva entirely The holotype was thought to be housed in the membranous; secondary gonopore placed at base collection of NKMU but we could not locate it of vesica in repose; sclerotized part of ductus sem- and we consider it as lost. inis twice as long as wide; vesica with two serrate We provide the following translation of the spicules, bearing toothlike outgrowths. type series translated from Chinese by our col- Female. Total length 4.4–5.2. COLORATION league Xiaojing Wang (personal commun., trans- (fig. 4): Head and labium. As in male. Antenna: lated from Hu and Zheng, 2001): Similar to male, rarely segment II pale brown basally. Thorax: Similar to male, but posterior Body slender, yellowish brown, lateral sides parallel part of pronotum uniformly yellow to pale brown to each other, with pale hairs; vertex yellowish or with brown humeral angles; mesoscutum and brown, with reddish tinge, with rare pale semierect scutellum yellow to pale brown, mesoscutum setae, ratio of distance between anterior margin of medially and scutellum basally sometimes head to posterior margin of eye and distance brown; thoracic pleura pale brown to brown. between posterior margin of eye to collar equal to Hemelytron: As in male. Legs: Coxae whitish yel- 0.3:0.4; frons almost flat, frons and lateral sides of low, femora whitish yellow basally and yellow head yellowish brown; base of labium brown; neck apically, hind femora rarely with pale brown yellowish brown, with slight reddish tinge; labium marking at apical half, rarely uniformly whitish yellowish brown, reaching posterior margin of fore- yellow; tibia and tarsi whitish yellow to yellow. coxae; antennae yellowish brown, apex of antennal Abdomen: Whitish yellow to yellow ventrally, segment I and apical half of antennal segment II segment IX pale brown or reddish, stripe later- slightly darkened, with rare, pale and semierect ally and dorsal surface reddish to reddish brown. hairs, the length of hairs on antennal segment I ca. SURFACE AND VESTITURE: As in male. 1.5× as long as antennal segment I width; hairs on STRUCTURE AND MEASUREMENTS: Struc- antennal segment II subequal to the antennal seg- ture as in male; body ca. 4.1–4.4× as long as pro- ment I diameter; pronotum yellowish brown, with notum width; vertex ca. 1.8–2.0× as wide as eye rare pale and semierect hairs, posterior margin diameter; antennal segment I ca. 1.3–1.5× as brown, areas near posterior margin with slightly long as head width, ca. 0.6–0.8× as long as pro- pale red tinge; lateral margins with slight depres- notum width; segment II ca. 1.8–2.2× as long as sion; scutellum yellowish brown, length of hairs ca. head width, ca. 0.9–1.1× as long as pronotum 2.0× as long as antennal segment I width; inner part width; pronotum ca. 1.1–1.3× as wide as long of clavus with the main part yellowish brown, outer and ca. 2.0–2.1× as wide as head. Genitalia (fig. part of clavus brown, with rare and pale semierect 16L, M): Dorsal labiate plate wider than hairs, equal in length to the hairs on scutellum; apodemes of second valvulae, with distinct stria- corium yellowish brown, semitransparent, with arc- tions, without membranous ridge medially; shaped, broad and dark brown marking, reaching semicircular sclerite and sclerotized rings absent; 1/5 of claval length; cuneus yellowish brown, ¼ apex lateral oviducts placed in posterior part, far from of inner margin red, length of outer margin twice as posterior margin; spermathecal gland attached long as base; membrane pale yellowish brown, pos- 74 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
terior half of wing cell and middle of membrane and Zheng, 2001: fig. 2b, c), which are very dis- brownish yellow, veins yellowish brown, or with tinct in comparison to all other Felisacus species. reddish tinge, apical angle round; legs yellowish Based on this, we treat those additional speci- brown, apex of femora and basal area of tibiae with mens as F. curvatus. red tinge; femora with rare, pale and semierect Felisacus curvatus and F. wangae are sister hairs, varying in length, short hairs subequal to species and have similar female genitalia with middle part of femora width, longest ca. 1.5× as distinct striations (cf. fig. 16L with 16N), the long as femur width; tibiae with hairs more right-hand side of the genital capsule is slightly adpressed than those on femora, hairs on half of folded (cf. figs. 14J and 15M), and the apical apex dense and uniform, short hairs subequal to sclerotized part of the ductus seminis is twice as tibiae width, longest setae is ca. 2.0× as long as tibiae long as wide (cf. fig. 10E with 10M). Felisacus width; prosternum yellowish brown, mesosternum wangae differs in that the marking on the inner brown, with middle part yellowish brown, with pale margin of the corium is narrow, and its posterior semierect hairs; males with ventral side of abdomen part not widened (fig. 7), the labium reaches the yellowish brown, lateral side of the second segment middle of the mesosternum, and in the shape of with one long, small and red marking, segments III– the aedeagal spicules (fig. 10M). VII with red lateral margins; genital segment yel- Material examined: CHINA: Yunnan: lowish brown; females with lateral sides of Ruili, 24.01655°N 97.85099°E, 1200 m, 29 Jul abdominal II–VII segments red. 2006, Fan, 1♂ (00017888) (NKMU). LAO PEO- PLE’S DEMOCRATIC REPUBLIC: Vientiane Measurements: Head length 0.6; head width 0.5; Prov.: Ban Van Eue, 17.96666°N 102.6°E, 14 Apr vertex width 0.3; antennal segments ratio 1965–15 Apr 1965, J.L. Gressitt, 1♀ (00043876) 1.2:1.5:1.8:1.5; pronotum length 1.0, pronotum (BPBM). VIETNAM: Ha Noi: 70 km NWW Ha width 1.0; outer margin of corium 1.7; outer length Noi BaVi, 21.087°N 105.302°E, 22 Nov 1990, of cuneus 0.7; base of cuneus width 0.3; length Belokobylskij, 1♀ (00017883) (ZISP). Ha Son between apex of cuneus and end of membrane 0.6; Binh: Hoa Binh: Near Mai Chau, 20.667°N body length 4.4–4.6. 105.084°E, 01 Nov 1990, Belokobylskij, 4♂ (00017882, 00271519), 2♀ (00017886) (ZISP); Type: Male, from Hainan, 1964 03 28, Shengli LIU. 02 Nov 1990, Belokobylskij, 3♂ (00017879– This species is similar to F. ochraceus in yellowish- 00017881), 2♀ (0001788400017885) (ZISP). brown coloration, and marking around near inner margin of corium. The latter species differs in apex Felisacus dauloi Woodward of antennal segment II not red; 2/3 of inner part of clavus yellowish brown, outer part of clavus brown; Figures 4, 8G, 11X, Y, 14K, 18 apical 1/6 of inner margin of cuneus pale red; the Felisacus dauloi Woodward, 1958: 236 (original band near inner margin of corium broader and lon- description). ger; pronotum with posterior margin and posterior Diagnosis: Recognized by the following half of lateral margin brown; parameres different; combination of characters: body mostly whitish ratio of antennal segments I–III in F. curvatus: yellow or yellow without brown markings, with 0.8:1.0:1.2 (in F. ochraceus 0.8:1.2:0.8: ratio between reddish tinge and yellow or red cuneus (fig. 4); length and width of cuneus in F. curvatus: 0.66: 0.27, antennal segment I cylindrical (as in Namyatova in F. ochraceus is 0.52:0.20). et al., 2016: fig. 8A), transverse depression on We have examined additional specimens from head extending present dorsally and laterally, southeast Asia, which fit the description of F. vertex upraised (as in Namyatova et al., 2016: fig. curvatus. They possess similar parameres (Hu 6D); relatively large body 3.8–4 in male and 4.3– 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 75
4.5 in female; cuneus ca. 3× as long as base; Corium smooth, with shallow and scarce punc- medial part of right paramere only slightly wider tures. Dorsum, antennal segment I and femora than basal part, shorter than basal and apical clothed with setae mostly as long as or some- parts combined; outer margin of right paramere what shorter than antennal segment II diameter; almost straight (fig. 11X); apical part of left para- abdomen clothed with suberect mostly short mere not widened, ca. 3× as long as wide (fig. simple setae. STRUCTURE AND MEASURE- 11Y); six spicules vesical spicules, including spic- MENTS: Body ca. 4.4–4.7× as long as pronotum ules A and B (fig. 8G). width. Head: Depression, delimiting occipital Redescription: Male. Total length 3.8–4.0. region, present dorsally and laterally (as in COLORATION (fig. 4): Head: Mostly yellow Namyatova et al., 2016: fig. 4E); distance between with reddish tinge and marking; longitudinal depression and pronotum distinctly shorter than sulcus sometimes yellow brown, tubercle around eye diameter; longitudinal sulcus on dorsal sur- antennal fossae, maxillary plate and buccula face of head longer than eye diameter; distance sometimes whitish yellow; clypeus yellow with from eye to pronotum as long as eye diameter, longitudinal red stripe; mandibular plate red or not swollen laterally (as in Namyatova et al., yellow with reddish marking. Eye brown to dark 2016: fig. 4E); vertex ca. 1.4–1.7× as wide as eye; brown, often with reddish margins. Labium: Yel- vertex raised (as in Namyatova et al., 2016: fig. low, segment III with pale brown or red stripe 6D); buccula ca. 0.2–0.25× as long as clypeus. ventrally. Antenna: Segment I yellow to reddish Labium: Almost reaching posterior part of brown, darkened apically; segment II reddish mesosternum; segments I and II strongly brown. Thorax: Pronotum yellow with pale reduced, combined shorter than half of segment brown anterior margin, often with longitudinal III; segment I shorter than wide (as in Namya- red stripe laterally; mesoscutum yellow; scutel- tova et al., 2016: figs. 6E, 9C); segment II slightly lum yellow, sometimes with red longitudinal longer than wide, its dorsal surface elongate stripe; thoracic pleura yellow with reddish areas, posteriorly; segment III ca. as long as ventral sometimes mostly red; scent gland evaporative side of head length; segment IV ca. 1.5× as long area whitish often with reddish tinge apically. as segment III. Antenna: Segment I cylindrical Hemelytron: Mostly colorless and translucent; (as in Namyatova et al., 2016: fig. 8A), ca. 1.7– inner part of clavus sometimes opaque, whitish 1.8× as long as head width, ca. 1.1× as long as yellow to yellow, with margins yellow to pale pronotum width; segment II ca. 2.2× as long as brown or reddish; corium with area along inner width of head, ca. 1.4× as long as width of pro- margin of corium pale brown or red, narrow notum. Thorax: Anterior part of pronotum and short, not reaching R+M; embolium some- slightly shorter than posterior part; collar delim- times yellow, with yellow to pale brown or red ited; posterior part of pronotum slightly apex and margins; cuneus with margins and upraised; posterior margin of pronotum con- often apex yellow or red, sometimes mostly yel- cave; pronotum ca. 1.2–1.3× as wide as long and low; membrane with grayish tinge, with mem- ca. 1.4–1.7× as wide as head; mesoscutum brane cell yellow to pale brown or red. Legs: slightly exposed. Hemelytron: Area along inner Coxae whitish yellow; femora whitish yellow margin of corium flat; inner margin of cuneus basally and yellow to red apically; tibiae whitish convex (as in Namyatova et al., 2016: fig. 13E), yellow to yellow, yellow or reddish basally; tarsi medial margin of cuneus ca. 3× as long as base. pale brown, sometimes tarsal segment I whitish Genitalia: Genital capsule (fig. 14K) slightly lon- yellow. Abdomen: Ventral and lateral sides of ger than wide; ventral wall twice as long as dor- pregenital segments whitish yellow, dorsal sur- sal wall, with posterior margin of ventral wall face of pregenital segments and entire genital semioval, smooth, without outgrowth(s), its capsule red. SURFACE AND VESTITURE: apex inclined slightly leftward, not curved; sides 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 of genital capsule not modified; right margin of segment III slightly longer than segment II; seg- paramere socket angulate, left margin rounded; ment IV ca. 0.3× as long as segment III; prono- distance between paramere sockets ca. 0.3× as tum ca. 1.2–1.3× as wide as long and ca. 1.6–1.8× long as base of genital capsule. Right paramere as wide as head. Genitalia (as in Namyatova et (fig. 11X) distinctly curved in apical half; apex al., 2016: fig. 23F, G): Dorsal labiate plate wider straight dorsally; medial part only slighter wider than distance between apodemes of second val- than basal part, bearing setae, with outer margin vula; mostly smooth, without distinct striations, straight and inner margin only slightly convex; with semicircular sclerite and distinct sclerotized outer angle distinct; inner angle rounded, with- rings laterally; lateral oviducts placed almost out setae; basal part ca. 0.15–0.2× as long as rest medially, very close to each other, spermathecal of paramere. Left paramere (fig. 11Y) L-shaped; gland placed between lateral oviducts; dorsal apical part not flattened, with toothlike out- labiate plate with distinct tubercles, without growth on posterior side medially (as in fig. membranous lobe medially. 11G) and without outgrowth on dorsal surface; Distribution: Papua New Guinea (fig. 18). middle part widened, without swelling or out- Host plant: Ferns in wet tropics (Wood- growth; setae only on middle part near outer ward, 1958). margin. Aedeagus conjunctiva weakly sclero- Remarks: Antennal segments III–IV in males tized, secondary gonopore placed at base of were lost, in females only parts of antennal seg- vesica in repose; sclerotized part of ductus semi- ment IV remain. nis around secondary gonopore shorter than Discussion: Felisacus dauloi was described wide; vesica with six spicules, including spicules by Woodward (1958) from a single female. A and B (fig. 8G). One of us (A.A.N.) examined the holotype, Female. Total length 4.3–4.5. COLORATION preserved in the Queensland Museum. We (fig. 4): Head: Similar to male, sometimes pale have some other nontype specimens from brown posteriorly; mandibular plate yellow, yel- BPBM, collected also in Papua New Guinea, low with reddish tinge or red. Labium: As in and we found that they are very similar to the male. Antenna: Similar to male, segment I uni- type of F. dauloi and treat them as belonging formly yellow or pale brown, yellow at base, seg- to that species. ments III–IV brown to dark brown. Thorax: Felisacus dauloi is similar to F. filicicola, F. Similar to male, but mesoscutum and scutellum ochraceus, and F. tanna in coloration and exter- yellow, thoracic pleura sometimes with pale nal morphology (figs. 4–7).Felisacus filicicola brown markings. Hemelytron: As in male. Legs: differs from F. dauloi in the mostly yellow or Similar to male, tarsi pale brown, sometimes seg- reddish-yellow cuneus and the presence of ment I and II whitish to yellow. Abdomen: Ven- seven vesical spicules, including spicules A, B, tral side and partly lateral side of pregenital E, and G (fig. 8I). Felisacus ochraceus differs in segments and segment VIII whitish yellow, upper the cuneus uniformly yellow or with yellow or part of lateral side and dorsal surface of segments red inner part, the medial part of the right para- II–VIII and entire segment IX red. SURFACE mere is as wide as the basal part, and shorter AND VESTITURE: As in male. STRUCTURE than the basal and apical parts combined, the AND MEASUREMENTS: Structure as in male; outer margin of the right paramere is distinctly body ca. 4.2–4.6× as long as pronotum width; concave (fig. 13C), and there are five vesical vertex ca. 1.6–1.8× as wide as eye diameter; spicules, including spicules A, B, C, and H (fig. antennal segment I ca. 1.5–1.8× as long as head 9F). Felisacus tanna differs from F. dauloi in the width, ca. 0.9–1.1× as long as pronotum width; apex of the cuneus colorless often with a yellow segment II ca. 1.9–2.2× as long as head width, ca. tinge and is never red, the middle part of the 1.2–1.4× as long as pronotum width; antennal right paramere is broad, twice as wide as the 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 77 basal part (fig. 13T), and the vesica has five cal part of left paramere not widened, ca. 3× as spicules, including spicules A, B, and E (fig. long as wide (fig. 11AA); vesica with five spic- 9M). ules, including spicule B (fig. 8H). Material examined: Holotype: PAPUA Redescription: Male. Total length 4.0–4.2. NEW GUINEA: Eastern Highlands: Daulo COLORATION (fig. 5): Head: Whitish yellow to Pass, 20 Aug 1956–22 Aug 1956, T.E. Wood- pale brown, rarely with greenish tinge, often dor- ward, 1♀ (00201839) (QM). Additional mate- sal surface and sometimes lateral side and frons rial: PAPUA NEW GUINEA: Morobe Province: somewhat darker than rest of head; head often Kaisenik Rd, Wau Subdistrict, 7.38012°S with brown, dark brown, or red markings; clyp- 146.77817°E, 1100 m, 02 Mar 1978, W.C. eus whitish yellow to pale brown, often brown to Gagne, Pteris moluccanus (Pteridaceae), 4♀ dark brown apically. Eye dark brown, sometimes (00042366, 00042367, 00042277, 00042278) with reddish tinge, rarely eyes pale brown. Pteris moluccana (Pteridaceae), 3♂ (00042364, Labium: Whitish yellow to yellow, segments I–II 00042365, 00043882), 1♀ (00043881) (BPBM). sometimes with greenish tinge, segment III Northern Province: 9–10 km WSW of Tufi, SE sometimes with reddish tinge ventrally. Antenna: Cape Nelson, 9.1°S 149.3°E, 585 m, 08 Sep Segment I pale brown to brown with yellow base, 1982–13 Sep 1982, G.A. Samuelson, 1♂ rarely yellow with reddish tinge; segment II pale (00043880) (BPBM). brown to dark brown, rarely with reddish tinge; segments III–IV brown to dark brown. Thorax: Felisacus elegantulus (Reuter) Pronotum whitish yellow to yellow, rarely with reddish tinge; anterior margin of pronotum Figures 5, 8H, 11Z, AA, 18 sometimes pale brown; humeral angles of prono- Hyaloscytus elegantulus Reuter, 1904: 2 (original tum often pale brown to dark brown; pronotum description). sometimes uniformly colored; mesoscutum and Hyaloscytus elegantulus var. filicicola Kirkaldy, scutellum whitish to yellow, scutellum some- 1908: 377 (original description); Knight, times brown apically, often paler than mesoscu- 1935: 200 (revised status). tum; punctures between mesoscutum and Hyaloscytus elegantulus var. javanus Reuter, scutellum pale brown; thoracic pleura whitish 1908: 187 (original description; synony- yellow to yellow, rarely with greenish tinge; scent mized by Poppius, 1914: 148 with Felisacus gland evaporative area uniformly whitish, rarely glabratus (Motschulsky, 1863). with greenish tinge. Hemelytron: Mostly translu- cent, colorless; inner part of clavus sometimes Diagnosis: Recognized by the following opaque, and yellow with margins yellow to combination of characters: dorsum mostly yel- brown; corium with marking along inner margin low to pale brown, without reddish tinge or any of corium yellow to brown; embolium some- red markings; humeral angles of pronotum often times opaque and yellow, with pale brown to brown to dark brown; antennal segment I cylin- brown margins; cuneus with yellow to brown drical (as in Namyatova et al., 2016: fig. 8A); margins, outer margin often somewhat darker transverse depression on head extending later- than inner margin; membrane often with yellow ally; labium reaching posterior margin of meso- or grayish tinge; membrane cell yellow to pale sternum; dorsal surface of labial segment II brown. Legs: Coxae whitish yellow to yellow; elongate posteriorly (as in Namyatova et al., femora whitish yellow to yellow basally and 2016: fig. 6D); cuneus ca. 3× as long as base; darker, yellow to pale brown, apically, sometimes medial part of right paramere as long as apical with reddish tinge; tibia whitish yellow to pale part, only slightly wider than basal part; outer brown, often with paler, whitish to yellow, apical margin of right paramere straight (fig. 11Z); api- part, sometimes with reddish stripe; tarsi yellow 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 to pale brown, sometimes tarsal segment I paler, of abdomen. Genitalia: Genital capsule (Namya- whitish yellow to yellow. Abdomen: Whitish to tova et al., 2016: fig. 22L) ca. 1.5× as long as yellow, sometimes with greenish tinge. SUR- wide; ventral wall ca. 1.5× as long as dorsal wall, FACE AND VESTITURE: Corium smooth, with with posterior margin semioval, smooth, without shallow punctures. Dorsum, antennal segment outgrowth, slightly pointed, its apex slightly II, and femora clothed with setae as long as or inclined leftward, not curved; sides of genital shorter than antennal segment II diameter; capsule not modified; right paramere socket abdomen clothed with short erect setae, dis- slightly pointed, left paramere socket rounded; tinctly shorter than width of antennal segment distance between paramere sockets subequal to II. STRUCTURE AND MEASUREMENTS: half of genital capsule width at base. Right para- Body ca. 4.2–4.4× as long as pronotum width. mere (fig. 11Z) distinctly curved apically; apex Head: Depression delimiting occipital region straight; medial part only slighter wider than present dorsally and laterally (as in Namyatova et basal part, bearing setae, with outer margin al., 2016: fig. 4E); distance between depression straight or slightly concave and inner margin and pronotum distinctly shorter than eye diam- convex; outer angle distinct, slightly swollen; eter; longitudinal sulcus on dorsal surface of inner angle absent, without setae; basal part ca. head longer than eye diameter; distance from eye 0.25× as long as rest of paramere. Left paramere to pronotum distinctly longer than eye diameter, (fig. 11AA) L-shaped; apical part not flattened, not swollen laterally (as in Namyatova et al., with toothlike outgrowth on posterior side medi- 2016: fig. 4E); vertex ca. 1.7–1.9× as wide as eye, ally and without outgrowth on dorsal surface; upraised (as in Namyatova et al., 2016: fig. 6D); middle part widened, without swelling or buccula ca. 0.2–0.25× as long as clypeus. Labium outgrowth(s); setae only on middle part near (as in Namyatova et al., 2016: figs. 6D, 9C): outer margin. Aedeagus (Namyatova et al., 2016: Reaching posterior margin of mesosternum or fig. 22I) conjunctiva weakly sclerotized; second- slightly surpassing it; segments I and II strongly ary gonopore placed at base of vesica in repose; shortened, combined shorter than half of seg- sclerotization of ductus seminis around second- ment III; segment I shorter than wide; segment ary gonopore shorter than wide; vesica with five II as long as wide, its dorsal surface elongate pos- spicules, including small spicule B and unique teriorly; segment IV ca. 1.5× as long as segment large bifurcate spicule ventrally (fig. 8H). III. Antenna: Segment I cylindrical (as in Namy- Female. Total length 4.1–4.7. COLORATION atova et al., 2016: fig. 8A), ca. 1.5–1.8× as long as (fig. 5). Similar to male, rarely coloration of body, head width, ca. 0.9–1.0× as long as pronotum pronotum, thoracic pleura and inner part of cla- width; segment II ca. 1.9–2.1× as long as width vus pale brown, abdomen whitish to pale brown, of head, ca. 1.1–1.2× as long as pronotum width; rarely with greenish tinge, sometimes reddish segment III slightly longer than segment II; seg- apically. SURFACE AND VESTITURE: As in ment IV subequal to half of segment III. Thorax: male. STRUCTURE AND MEASUREMENTS: Anterior part of pronotum only slightly shorter Structure as in male; body ca. 4.2–4.3× as long as than posterior part; collar delimited; posterior width of pronotum; vertex ca. 1.8–2.1× as wide part slightly upraised; posterior margin concave; as eye diameter; antennal segment I ca. 1.4–1.7× pronotum ca. 1.3–1.4× as wide as long and ca. as long as head width, ca. 0.8–0.9× as long as 1.7–1.9× as wide as head; mesoscutum exposed. pronotum width; segment II ca. 1.9–2.1× as long Hemelytron: Area along inner margin of corium as head width, ca. 1.1–1.2× as long as pronotum almost flat; inner margin of cuneus convex (as in width; pronotum ca. 1.3–1.4× as wide as long Namyatova et al., 2016: fig. 13E), outer margin of and ca. 1.7–1.8× as wide as head. Genitalia (as in cuneus ca. 3× as long as base. Abdomen: Genital Namyatova et al., 2016: fig. 23F, G): Dorsal labi- capsule rotated left at right angle relative to rest ate plate wider than distance between apodemes 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 79 of second valvula; mostly smooth, without dis- paramere is broad, twice as long as wide (fig. tinct striations, with semicircular sclerite and 11Q), and there are four vesical spicules, includ- distinct sclerotized rings laterally; lateral ovi- ing spicules A and B (fig. 8F). ducts placed almost medially, very close to each Felisacus lordhowensis differs from F. elegantu- other, spermathecal gland placed between lateral lus in the labium reaching the posterior margin oviducts; dorsal labiate plate with distinct tuber- of the metasternum, the humeral angles of the cles, without membranous lobe medially. pronotum is whitish yellow to yellow, the prono- Distribution: Australia, New Zealand (fig. 18). tum is brown posteriorly, and there are six vesi- Host plants: The species was collected mostly cal spicules, including spicules B and E (fig. 8R). from ferns and a few specimens were also col- Felisacus schuhi differs from F. elegantulus in lected from angiosperms. The host plant for F. the humeral angles often yellow to pale brown, elegantulus is primarily Pteris tremula R.Br. (Pteri- the inner part of the clavus often brown, the daceae) in New Zealand (35 specimens). Wood- labium only slightly surpassing the middle of the ward (1954) also reported that many specimens of mesosternum, and there are five spicules, includ- F. elegantulus were collected from Hypolepis muel- ing spicules A, B, and C (fig. 9J). leri N.A. Wakef. (Dennstaedtiaceae) in New South Material examined: AUSTRALIA: New Wales, Australia (5 specimens); Pteris sp. (Pterida- South Wales: Bola Creek, Royal National Park, ceae) in New Zealand (2 specimens); and Polysti- 34.11495°S 151.06544°E, 27 Feb 1989, G. Cas- chum sp. (Dryopteridaceae) in New Zealand (35 sis, 2♂ (00017724, 00017725), 1♀ (00017735) specimens). Plant records among angiosperms are (AM). Jamieson Park, Narrabeen, 33.7169°S considered sitting records only, and include 151.28207°E, 01 Feb 1996, B. Day and P. Serov, Acmena smithii (Poir.) Merr. (Myrtaceae) in New 3♀ (00017739–00017741) (AM). Kings Rd., South Wales (1 female specimen); Dendrocnide Federal, 28.65331°S 153.43432°E, 02 Jan 2012, excelsa (Wedd.) Chew (Urticaceae) in Queensland G. Cassis, 10♂ (00400524, 00400110– (1 female specimen); and Pomaderris apetala 00400116, 00400524, 00191459), 11♀ Labill. (Rhamnaceae) in Tasmania (2 specimens). (00191458, 00191460, 00400501, 00400118– Discussion: Felisacus elegantulus was 00400123, 00400501, 00191458), 2 sex described by Reuter (1904) from two males and unknown (00191460, 00400117), 10 juvenile we could not locate the syntypes, which we pre- (00400124–00400133) (UNSW). Minnamurra sume are lost. Nevertheless, there are abundant Falls nr Kiama, 34.63333°S 150.71666°E, 07 Jan specimens, collected throughout Australia and 1961, D.K. McAlpine, 1♂ (00017552) (AM). New Zealand, fitting the original description. Mount Keira, 34.24°S 150.51°E, 09 May 1970, This species has been treated as F. elegantulus by D.K. McAlpine and G.A. Holloway, 1♀ Eyles and Schuh (2003), and we also consider it (00017738) (AM). Porter’s Dam Road, 16 km as conspecific toF. elegantulus. There is another NM of Milton, 35.17925°S 150.44048°E, 09 Feb species of Felisacus, collected in Queensland, but 1974, Z. Liepa, 2♂ (00033774, 00033775) it has different coloration from the species, (ANIC). Rotary L/out, Pigeon House Ra NW described by Reuter (1904) (see F. australicus). of Milton, 35.35061°S 150.26785°E, 22 Feb See generic discussion for the explanation of 1979, Z. Liepa, 1♀ (00033771) (ANIC). Royal nomenclatural confusion in literature. National Park, Warumbul Picnic Area, Felisacus elegantulus is similar to F. caledoni- 34.06667°S 151.1048°E, 20 m, 14 Nov 2001, cus, F. lordhowensis, and F. schuhi in the color- Cassis, Schuh, Schwartz, Silveira, Hypolepis ation and the structure. Felisacus caledonicus muelleri N.A. Wakef. (Dennstaedtiaceae), det. differs in the lateral margins of the collar being NSW staff NSW666413, 3♂ (00021377, often brown, the humeral angles of the prono- 00021375, 00021376), 2♀ (00021373, tum are whitish yellow, the apical part of the left 00021374) (AM). S Coast of Durras Lake, 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
35.65613°S 150.27268°E, 22 Feb 1965, D.H. 36.84291°S 174.73087°E, 30 Dec 1997, G. Cas- Colless, 2♀ (00033769, 00033770), 1 sex sis, Polystichum sp. (Dryopteridaceae), 20♂ unknown (00033772), 2♂ (00033777, (00017562, 00017563, 00017742–00017745, 00033778) (ANIC). Sydney Lea, 33.87309°S 00017783–00017796), 1 juvenile (00017559), 151.16901°E, no date provided, collector 14♀ (00017797–00017810) (AM). Upokorau unknown, 1♂ (00017723) (AM). Wallaga Lake, Valley, 15 km W Takou Bay, Northland, Bermagui, 36.42279°S 150.06441°E, 21 Jul 35.13883°S 173.84274°E, 16 Jan 1998, G. Cas- 1973, Z. Liepa, 3♀ (00033766–00033768), 1♂ sis, 10♂ (00017746–00017755), 2 sex unknown (00033776) (ANIC). Wentworth Falls, Blue (00017560, 00017561), 7 juvenile (00017756– Mountains, 33.71083°S 150.37497°E, 20 Nov 00017762), 20♀ (00017763–00017782) (AM) 1958, D.K. McAlpine, 1♀ (00017557) (AM). North Auckland: Auckland, 36.8667°S approx. 24 km NNW of Gloucester, 32.048°S 174.7667°E, 14 Mar 1996, Tomkovich, 1♂ 151.903°E, 05 Dec 1990, G. and T. Williams, (00271527) (ZISP). Wellington: Whirokino Acmena smithii (Poir.) Merr. and L.M. Perry [Whirikino], Manuwatu, 40.51826°S (Myrtaceae), 1♀ (00017737) (AM). 175.25021°E, 7 m, 07 Jan 1953, T.E. Wood- Queensland: 5 mi E Paluma, Northern ward, Pteris sp. (Pteridaceae), 1♂ (00018975), Queensland, 19.00392°S 146.24205°E, 18 Jan 1♀ (00018974) (ISNB); 27 Jan 1957, T.E. 1970, G.A. Holloway, 1♀ (00017558) (AM). Woodward, 1♀ (00021557) (AMNH). Conondale Range, Bundaroo Creek, 26.683°S 152.617°E, 03 Dec 1985, G. Cassis and D. Felisacus fedori, sp. nov. Bickel, 5♀ (00017726–00017730), 4 juvenile (00017731–00017734), 8♂ (00017663– Figures 4, 10G, 11AB, AC, 14L, 18 00017667, 00017720–00017722) (AM). Mt. Diagnosis: Recognized by the following Edith Forest road, 1.5mi off Danbulla Road, combination of characters: marking along inner 17.1°S 145.633°E, 06 May 1967, D.H. Colless, margin of corium distinct, brown, its anterior 1♀ (00034248) (ANIC). Mt. Glorious, 27.333°S part widened, not reaching R+M (fig. 4), poste- 152.767°E, 16 Feb 1961–20 Feb 1961, L. and M. rior part, reaching R+M; posterior part of prono- Gressitt, 1♂ (00042309) (BPBM). Mt. Hypipa- tum mostly yellow, brown to dark brown only mee, National Park “The Crater,” 17.42388°S along posterior margin; cuneus red; inner mar- 145.48388°E, 25 Apr 1998, G. Cassis, Dendroc- gin of medial part of right paramere convex, nide excels (Urticaceae), 1♀ (00017736) (AM). without swelling; inner angle of right paramere Pinnock [Pinock] R., Gorge Hogback Ra., with short and acute outgrowth (fig. 11AB); WSW of Bundaberg, 25.03481°S 151.80731°E, medial part of left paramere with wide outgrowth 23 Sep 1972, H. Frauca, 2♂ (00033779, (fig. 11AC); distal half of ductus seminis sclero- 00033780) (ANIC). Woombye, nr. Nambour, tized, sickle shaped apically; vesica without spic- 26.665°S 152.976°E, 11 Oct 1965–16 Oct 1965, ules (fig. 10G). D.H. Colless, 2♀ (00033764, 00033765), 1♂ Description: Male. Total length 4.4–4.5. (00033773) (ANIC). Tasmania: Mt. Field COLORATION (fig. 4): Head: Mostly yellow with National Park, Russell Falls Visitor Centre, red stripes on frons above antennal fossa and 42.68151°S 146.7168°E, 167 m, 16 Jan 2004, sometimes laterally behind eye and with reddish M.D. Schwartz and P.P. Tinerella, Pomaderris markings on tubercle of antennal fossa; ventral apetala Labill. (Rhamnaceae), det. NSW staff side of head and buccula whitish yellow. Eye dark NSW658218, 1♂ (00021542) (AMNH). Victo- brown with reddish tinge. Labium: Uniformly ria: [Victoria], no date provided, Unknown, whitish yellow or with segments II–IV yellow. 2♀ (00005120) (MVMA). NEW ZEALAND: Antenna: Segment I yellow with reddish tinge; Auckland: Cremorne St, Herne Bay, Auckland, segment II reddish brown; segments III–IV 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 81 brown. Thorax: Pronotum yellow, its posterior surpassing prosternum; each of segments I and II part slightly paler than anterior part, with brown subequal to half of segment III; dorsal surface of humeral angles and medial stripe or all those labial segment II not elongate (as in Namyatova markings merged; mesoscutum brown; scutellum et al., 2016: fig. 6E); segment III shorter than ven- yellow to pale brown with brown medial stripe, tral side of head; segment IV ca. 1.5× as long as this stripe darker basally and pale apically; tho- segment III. Antenna: Segment I widened medi- racic pleura mostly pale brown with dark brown ally (as in Namyatova et al., 2016: fig. 8B), ca. markings; scent gland evaporative area whitish 1.1–1.2× as long as head width, ca. 0.9× as long yellow. Hemelytron: Mostly translucent; inner as pronotum width; segment II 2.0–2.2× as long part of clavus opaque, brown; outer part of clavus as head width, 1.3–1.4× as long as pronotum pale brown, dark brown basally; corium colorless width; antennal segment III almost as long as seg- with brown marking along inner margin of ment II; segment IV slightly shorter than segment corium, its anterior part widened, not reaching III. Thorax: Anterior part of pronotum shorter R+M vein, posterior part reaching R+M vein; than posterior part; collar delimited; posterior embolium colorless or pale brown, with brown part slightly upraised, posterior margin of prono- base and outer margin and reddish apex; cuneus tum slightly concave; pronotum ca. 1.2× as wide opaque, uniformly red; membrane brown basally as long and 1.6× as wide as head; mesoscutum and whitish and with greyish tinge apically; slightly exposed or not exposed. Hemelytron: membrane cell pale brown to brown. Legs: Coxae Area along inner margin of corium swollen; inner whitish yellow; femora whitish yellow basally and margin of cuneus straight, outer margin of cuneus yellow apically, forefemur with reddish stripe api- 2× longer than width at base. Abdomen: Genital cally and medially, hind femur with red or brown capsule rotated 45° relative to other segments. marking in apical half; tibiae yellow basally and Genitalia: Genital capsule (fig. 14L) ventral wall whitish yellow apically, with red stripe distinct ca. 1.5× as long as dorsal wall, with posterior basally and pale apically; tarsi whitish yellow. margin of ventral wall not curved dorsally, with Abdomen: Whitish yellow, with brown dorsum small tooth-shaped outgrowth; right side of geni- and genital capsule. SURFACE AND VESTI- tal capsule curved, left side of genital capsule not TURE: Corium smooth, with shallow and scarce widened; right paramere sockets angulate, left punctation. Dorsum with setae subequal to paramere socket rounded; distance between para- antennal segment II diameter; antennal segment mere sockets subequal to 1/4 of genital capsule I clothed with suberect setae shorter than anten- base width. Right paramere (fig. 11AB) distinctly nal segment II diameter; femora with suberect curved in apical half; apex concave posteriorly; setae subequal to or longer than antennal seg- medial part narrower than basal part, outer mar- ment II diameter; abdomen clothed with short gin of medial part convex and inner margin con- erect setae. STRUCTURE AND MEASURE- cave, without swelling, without setae; outer angle MENTS: Body ca. 4.0–4.1× as long as pronotum absent; inner angle present, with straight out- width. Head: Depression delimiting occipital growth, bearing setae; basal part of right param- region present only dorsally, not extending later- ere ca. 0.3× as long as rest of paramere. Left ally (Namyatova et al., 2016: fig. 6E); distance paramere (fig. 11AC) Apical part not flattened, between depression and pronotum slightly with tooth on posterior side apically and with shorter than eye diameter; longitudinal sulcus on outgrowth on dorsal surface; middle part with dorsal surface of head shorter than eye diameter; short and wide outgrowth bearing setae. Aedeagus distance from eye to pronotum subequal to eye (fig. 10G) vesica without spicules; distal half of diameter, not swollen laterally; vertex ca. 1.5–1.6× ductus seminis sclerotized, secondary gonopore as wide as eye diameter; vertex raised; buccula ca. placed near phallotheca mouth in repose; ductus 0.3× as long as clypeus height. Labium: Slightly seminis hooked apically. 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
Female. Total length 4.9–5.9. COLOR- gins, and the outgrowth on the inner part of ATION (fig. 4): Head: Similar to male, but right paramere abrupt (fig. 11T). without red stripes and markings; labial seg- Material examined: Holotype: INDONE- ments II–IV often yellow, segment II some- SIA: Sulawesi Utara: Dumoga-Bone National times brown ventrally; antennal segment I Park, Plot C, 400 m, 11 Feb 1985, Royal Ento- sometimes reddish brown apically. Thorax: mological Society of London: Project Wallace, Similar to male; collar often brown to dark 1♂ (00019529) (BMNH). Paratypes: INDONE- brown; scutellum yellow to pale brown with SIA: Sulawesi Utara: Dumoga-Bone National brown stripe medially; mesoscutum yellow, Park, Gunung Ambang Forest Reserve, near brown basally or uniformly brown; thoracic Kotamobagu, Fog. 7, 0.55°N 123.98333°E, 1200 pleura yellow to pale brown with dark brown m, 18 Feb 1985, Royal Entomological Society metepimeron, sometimes with markings also of London: Project Wallace, 2♀ (00019534, dark brown. Hemelytron: Similar to male, but 00019531) (BMNH). Dumoga-Bone National outer part of clavus sometimes brown, same Park, Plot B, Fog. 6/14, 315 m, 13 Feb 1985, color as inner part; embolium pale brown with Royal Entomological Society of London: Proj- brownish base and reddish apex. Legs: Similar ect Wallace, 1♀ (00019535) (BMNH). to male, but marking in apical half of femur red Dumoga-Bone National Park, plot B, Fog 3, or reddish brown and sometimes absent on 315 m, Feb 1985, Royal Entomological Society middle femur; abdomen similar to male, some- of London: Project Wallace, 1♀ (00019530), times uniformly whitish yellow. SURFACE 1♂ (00019528) (BMNH). AND VESTITURE: As in male. STRUCTURE AND MEASUREMENTS: Structure as in male, Felisacus filicicola (Kirkaldy) body ca. 3.8–4.0× as long as pronotum width; vertex ca. 1.1–1.5× as wide as eye diameter; Figures 5, 8I, 11AD, AE, 14N, 18 antennal segment I ca. 1.2–1.3× as long as head Hyaloscytus elegantulus var. filicicola Kirkaldy, width, ca. 0.8–1.0× as long as pronotum width; 1908: 377 (original description). segment II ca. 1.9–2.2× as long as head width, Felisacus filicicola Knight, 1935: 200 (comb. ca. 1.1–1.2× as long as pronotum width; prono- nov.). tum ca. 1.2–1.4× as wide as long and ca. 1.6– 1.9× as wide as head. Diagnosis: Recognized by the following Etymology: The species is named after Fedor combination of characters: coloration of dorsum V. Konstantinov (St. Petersburg State University, generally yellow, often with reddish tinge or Russia), who helped us to get many specimens markings; cuneus often yellow to red, or at least for this study. with reddish (fig. 4); cylindrical antennal seg- Distribution: Indonesia, Sulawesi Is. (fig. 18). ment I (as in Namyatova et al., 2016: fig. 8A), Host plants: Unknown. transverse depression delimiting occipital region Discussion: Antennal segments, and the extending laterally; vertex upraised (as in Namy- middle and hind legs are lost in the female; the atova et al., 2016: fig. 6D); body length in male female genitalia were not dissected. Felisacus 3.5–3.8 and in female 3.5–4.2; labium reaching fedori is similar to F. cristobalus in the marking posterior margin of mesosternum; medial part of along the inner margin of corium and anterior right paramere slightly wider than basal part, part, which is broad and not reaching R+M (fig. shorter than basal and apical parts combined; its 4); in contrast, the latter species differs from F. outer margin straight (fig. 11AD); apical part of fedori in the pronotum mostly brown to dark left paramere not widened, ca. 3× as long as wide brown with pale brown stripe medially, the (fig. 11AE); vesica with seven spicules, including cuneus translucent, colorless with brown mar- spicules A, B, E, and G (fig. 8I). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 83
Redescription: Male. Total length 3.5–3.8. lateral and ventral side red; dorsal surface of pre- COLORATION (fig. 5): Head: Yellow to pale genital segments often red, sometimes yellow with brown, somewhat darker dorsally, sometimes with red medial longitudinal stripe; genital capsule yel- reddish tinge and reddish markings; longitudinal low to pale brown, sometimes with reddish tinge. sulcus on dorsal surface yellow to pale brown; SURFACE AND VESTITURE: Corium smooth, clypeus often darkened apically rarely with red- often with very shallow, scarce punctures. Body dish longitudinal stripe. Eye brown to dark brown, clothed with pale simple setae; dorsum and fem- often with reddish tinge. Labium: Yellow to pale ora with setae subequal to or shorter than anten- brown, segment I and III sometimes somewhat nal segment II diameter; antennal segment I darker than other segments, segment III some- clothed with suberect setae mostly shorter than times with reddish tinge. Antenna: Segment I yel- antennal segment II diameter; abdomen clothed low to reddish or reddish brown, often paler at with suberect setae, subequal to or shorter than base and darkened apically; segment II yellow to width of antennal segment II. STRUCTURE AND pale brown with reddish tinge, often darker than MEASUREMENTS: Body ca. 4.2–4.4× as long as segment I; segments III–IV brown to dark brown. width of pronotum. Head: Depression delimiting Thorax: Pronotum yellow to pale brown, some- occipital region present dorsally and laterally (as times with reddish longitudinal stripe at sides; in Namyatova et al., 2016: fig. 4E); distance posterior part of pronotum yellow or pale brown, between depression and pronotum distinctly sometimes posterior margin brown laterally; shorter than eye diameter; longitudinal sulcus on mesoscutum yellow to pale brown; scutellum yel- dorsal surface of head longer than eye diameter; low to pale brown, often brown to dark apically, distance between eye and pronotum slightly lon- sometimes mostly pale brown, yellow to pale ger than eye diameter, not swollen laterally (as in brown basally; thoracic pleura yellow to pale Namyatova et al., 2016: fig. 4E); vertex ca. 1.6– brown; scent gland evaporative area whitish yel- 2.0× as wide as eye, upraised (as in Namyatova et low to yellow, often with yellow or red apex. Hem- al., 2016: fig. 6D); buccula 0.15–0.2× as long as elytron: Mostly translucent and colorless; inner clypeus height. Labium: Reaching posterior mar- part of clavus whitish yellow to pale brown, some- gin of mesosternum or slightly surpassing it; seg- times red or brown apically, sometimes opaque; ments I and II strongly reduced, combined shorter outer margin of clavus sometimes whitish yellow, than half of segment III; segment I shorter than sometimes with brown tinge or pale brown api- wide (as in Namyatova et al., 2016: figs. 6D, 9C); cally; marking along inner margin of corium yel- segment II slightly longer than wide, elongate dor- low to brown; embolium sometimes with yellow sally; segment III slightly longer than ventral side tinge, rarely with reddish margins and yellow or of head length; labial segment IV ca. 1.5× as long red apically; cuneus yellow to reddish yellow, as segment III. Antenna: Segment I cylindrical (as often with whitish yellow outer part, rarely cuneus in Namyatova et al., 2016: fig. 8A), ca. 1.6–1.8× as with yellow tinge and yellow margins; membrane long as head width, ca. 1.0–1.2× as long as prono- sometimes yellow or with grayish tinge, with yel- tum width; segment II ca. 1.9–2.1× as long as low to pale brown membrane cell. Legs: Coxae width of head, ca. 1.3–1.4× as long as pronotum whitish yellow to yellow; femora whitish yellow to width; segment III slightly longer than segment II, yellow, darkened apically, sometimes reddish api- segment IV ca. 0.3× as long as segment III. Tho- cally; tibiae yellow to pale brown, paler apically, rax: Anterior part of pronotum slightly shorter sometimes with reddish tinge basally, rarely uni- than posterior part; collar delimited, posterior formly colored; tarsi yellow to brown, often with part slightly upraised; pronotum ca. 1.2–1.3× as tarsal segment I whitish yellow to yellow. Abdo- wide as long and ca. 1.4–1.6× as wide as head with men: Ventral and lateral sides of pregenital seg- posterior margin concave; mesoscutum more or ments often whitish yellow to yellow, sometimes less exposed. Hemelytron: Area along inner mar- 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 gin of corium almost flat; inner margin of corium in male. STRUCTURE AND MEASUREMENTS: convex (as in Namyatova et al., 2016: fig. 13E), Structure as in male; body ca. 4.1–4.5× as long as outer margin of corium ca. 2.5–3× as long as base. pronotum width; vertex ca. 1.6–1.9× as wide as Abdomen: Genital capsule rotated left at right eye diameter; antennal segment I ca. 1.4–1.9× as angle relative to rest of abdomen. Genitalia: Geni- long as head width, ca. 0.9–1.3× as long as pro- tal capsule (fig. 14N) ca. 1.5× as long as wide; ven- notum width; segment II ca. 1.8–2.2× as long as tral wall ca. 1.5× as long as dorsal wall, with head width, ca. 1.2–1.4× as long as pronotum posterior margin of ventral wall semioval, smooth, width; pronotum ca. 1.2–1.3× as wide as long not curved, without outgrowth(s), its apex placed and ca. 1.4–1.7× as wide as head. Genitalia (as in medially; margins not modified; right paramere Namyatova et al., 2016: fig. 23F, G): Dorsal labi- socket slightly acute and left paramere socket ate plate wider than distance between apodemes rounded; distance between paramere sockets ca. of second valvula; mostly smooth, without dis- 0.7× as long as genital capsule broad at base. Right tinct striations, with semicircular sclerite and paramere (fig. 11AD) distinctly curved in apical distinct sclerotized rings laterally; lateral ovi- half, apex straight; medial part only slightly wider ducts placed almost medially, very close to each than basal part, bearing setae, with outer margin other, spermathecal gland placed between lateral straight and inner margin convex; outer angle dis- oviducts; dorsal labiate plate with distinct tuber- tinct, not swollen; inner angle rounded, without cles, without membranous lobe medially. setae; basal part of right paramere ca. 0.15–0.2× as Distribution: Fiji (Viti Levu Is., Ovalau Is., long as rest of paramere. Left paramere (fig. 11AE) Kioa Is., Qamea Is., Laucala Is., Taveuni Is.), L-shaped; apical part not flattened, with toothlike Western Samoa (Upolu Is.) and American Samoa outgrowth on posterior side medially (as in fig. (Tau Is.), Vanuatu (Efate Is.) (fig. 18). 11G) and without outgrowth on dorsal surface; Host plants: Brackenlike ferns (Kirkaldy, middle part widened, without swelling or out- 1908). growth; setae only on middle part near outer mar- Discussion: Felisacus filicicola was described gin. Aedeagus (general view as in Namyatova et by Kirkaldy (1908) from Fiji without noting the al., 2016: fig. 22I) vesica weakly sclerotized, sec- type deposition. According to Kirkaldy, this ondary gonopore placed at base of vesica in species is similar to F. elegantulus, but its color- repose; sclerotization of ductus seminis around ation is suffused with red. Kirkaldy’s types are secondary gonopore shorter than wide; vesica usually preserved in BPBM, BMNH, or CAS, with seven spicules, including spicules A, B, and but we did not find his filicicola material in the G (fig. 8I). BPBM nor in the BMNH. In addition, the web- Female. Total length 3.5–4.2. COLORATION site of CAS (http://research.calacademy.org/ (fig. 5): Head: Similar to male, rarely head uni- redirect?url = http://researcharchive.calacad- formly yellow with reddish tinge. Labium: Simi- emy.org/research/entomology/typesDB/default. lar to male, segments I–II sometimes with asp) does not refer to them. reddish tinge. Antenna: As in male. Thorax, legs, We have examined numerous specimens from and hemelytron: As in male. Abdomen: Ventral Fiji, Vanuatu, and Samoa that fit the description and lateral sides of pregenital segments and seg- of F. filicicola of Kirkaldy (1908). Felisacus filici- ment VIII often whitish yellow to yellow, some- cola is variable externally and can be confused times pale brown, sometimes lateral sides partly with a number of species, including F. albus, F. reddish, rarely entire lateral and ventral surfaces amboinae, F. dauloi, F. insularis, F. javanus, F. red; ventral and lateral sides of segment IX yel- linae, F. ochraceus, and F. solomonicus (figs. 4–7). low to reddish, sometimes pale brown, often All of them differ from F. filicicola by the shape darker than other segments; dorsal surface of and number of vesical spicules (figs. 8A, B, G, L, abdomen red. SURFACE AND VESTITURE: As M, P, 9F, L). Felisacus albus and F. insularis differ 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 85 in the medial part of the right paramere twice as 00042235), 1♀ (00042271) (BPBM). Waiyevo, wide as its basal part (figs. 11A, 12C). Felisacus Taveuni Is., 16.78411°S 179.98011°W, 50 m, Jan javanus and F. ochraceus differ in the medial part 1972, N.L.H. Krauss, 1♂ (00042345) (BPBM). of the right paramere being distinctly concave SAMOA: Upolu: Afiamalu, 13.91666°S (figs. 12G, 13C). 171.71666°W, 640 m, 06 Jan 1956, T.E. Wood- Material examined: AMERICAN SAMOA: ward, 1♂ (00037409) (QM). Malololelei, 13.9°S Manu’a Islands: E of Tau Village, Tau, Manua, 171.75°W, 610 m, 19 Jan 1956, R.A. Cumber, 1♂ 14.22929°S 169.51111°W, 160 m, 18 Feb 1965, (00037408), 2♀ 00037406, 00037407) (QM). Mt. G.A. Samuelson, 1♂ (00042335) (BPBM). Vaea, 13.85°S 171.75°W, 427 m, Sep 1969, N.L.H. Tavalogi Ridge, Tau Is., 14.23185°S 169.46567°W, Krauss, 1♀ 00042312) (BPBM). VANUATU: 160 m, 16 Feb 1965–19 Feb 1965, G.A. Samuel- Shefa: Efate Is.: Klem Butte, Efate Is., New Hebri- son, 1♀ (00042336) (BPBM). FIJI: Central Divi- des, 17.66666°S 168.26666°E, 175 m, 05 Dec 1978, sion: Bilo nr Lami, Viti Levu Is., 18.11092°S N.L.H. Krauss, 1♂ (00042229) (BPBM). Vila, 178.41011°E, 20 m, 27 Sep 1979, M.K. Kamath, Efate Is., 17.41836°S 168.34517°E, Aug 1950, S.N. Lal, G.A. and S.L. Samuelson, 2♂ (00042231, N.L.H. Krauss, 1♂ (00042242) (BPBM). Vila, 00042232) (BPBM). Colo-i-suva [Tholo-i-Suva], Efate Is., 0–100 m, 17.74097°S 168.32099°E, Mar Viti Levu Is., 18.02336°S 178.42032°E, Mar 1951, 1978, N.L.H. Krauss, 13♂ (00042238, 00042320– N.L.H. Krauss, 1♂ (00042346), 1 sex unkown 00042328, 00042330, 00042249, 00042226), 6♀ (00042347) (BPBM). Lami, Viti Levu Is., (00042256–00042261) (BPBM); Nov 1978, N.L.H. 18.11092°S 178.41011°E, Mar 1951, N.L.H. Krauss, Krauss Coll., 10♂ (00042244–00042246, 1♀ (00042270) (BPBM). Lami, Viti Levu Is., 00042239, 00042240, 00042316, 00042318; 18.11092°S 178.41011°E, 100 m, Jan 1979, N.L.H. 00042319, 00042331), 8♀ (00042220, 00042241, Krauss, 1♀ (00042269), 1♂ (00042230) (BPBM). 00042251–00042255, 00042221), 3 juvenile Levuka, Ovalau Is., 17.68039°S 178.82994°E, 100 (00042222, 00042225, 00042243) (BPBM); Dec m, Dec 1978, N.L.H. Krauss, 5♀ (00042359– 1978, N.L.H. Krauss, 2♂ (00042248, 00042227), 00042363), 2♂ (00042353, 00042354) (BPBM). 4♀ (00042262–00042265), 1 sex unknown Levuka, Ovalau Is., 17.68039°S 178.82994°E, 20 (00042267) (BPBM); Feb 1981, N.L.H. Krauss, 3♀ Oct 1924, E.H. Bryan, Jr., 6♂ (00042333, (00042223, 00042224, 00042266), 2♂ (00042228, 00042334, 00042348–00042351), 3♀ (00042355– 00042247) (BPBM), 3♂ (00018424, 00018425, 00042357) (BPBM). Northern Division: Dreketi- 00018429), 3♀ (00018426–00018428) (MZH). lailai, Vanua Levu Is., 16.56009°S 178.92191°E, 50 m, 12 Oct 1979, S.N. Lal, G.A. and S.L. Samuel- Felisacus glabratus (Motschulsky), status nov. son, 2♂ (00042344, 00042343), 1♀ (00042332) (BPBM). Nadilo, Qamea Is., 16.79355°S Figures 5, 8J, 11AF, AG, 14N, 18 179.7641°W, 25 m, 03 Oct 1979, M.K. Kamath, Liocoris glabratus Motschulsky, 1863: 87 (origi- S.N. Lal, G.A. and S.L. Samuelson, 2♀ (00042275, nal description). 00042276), 3♂ (00042237, 00042250, 00042342) Felisacus glabratus Distant, 1904: 439 (comb. (BPBM). Qilo Bay, Laucala Is., 16.75841°S nov.). 179.68381°W, 25 m, 03 Oct 1979, M.K. Kamath, Hyaloscytus elegantulus var. javanus Reuter, 1908 S.N. Lal, G.A. and S.L. Samuelson, 1♂ (00042236), (original description; synonymized by Pop- ♀ 1 (00042358) (BPBM). Savusavu, Vanua Levu pius, 1914: 148); status nov. Is., 16.77786°S 179.33976°E, 100 m, Mar 1978, N.L.H. Krauss, 2♀ (00042272, 00042273) Diagnosis: Recognized by the following (BPBM). S coast to center, Kioa Is., 16.65143°S combination of characters: mostly whitish yellow 179.91088°E, 30 m, 04 Oct 1979, M.K. Kamath, to yellow coloration of dorsum, cuneus colorless S.N. Lal, G.A and S.L. Samuelson, 3♂ (00042233– or with yellow tinge, translucent (fig. 4); antennal 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 segment I cylindrical (as in Namyatova et al., metasternum; segments I and II strongly 2016: fig. 8A); transverse depression delimiting reduced, combined shorter than half of segment occipital region on head extending laterally, ver- III; segment I shorter than wide (as in Namya- tex in male ca. 1.9× as long as eye diameter tova et al., 2016: figs. 6D, 9C); segment II slightly upraised (as in Namyatova et al., 2016: fig. 6D); longer than wide, elongate dorsally; segment III antennal segment I ca. 1.3× as long as head slightly longer than ventral side of head length; width; cuneus ca. 3× as long as wide; labium segment IV ca. 1.5× as long as segment III. Tho- reaching middle of mesosternum; ventral wall of rax: Anterior part of pronotum slightly shorter genital capsule ca. 1.5× as long as dorsal wall (fig. than posterior part; collar delimited; posterior 14N); medial part of right paramere twice as part of pronotum slightly upraised; posterior wide as basal part, and shorter than apical and margin of pronotum concave; pronotum ca. basal parts combined; its outer margin concave 1.3× as wide as long and ca. 1.6× as wide as and inner margin convex (fig. 11AF); vesica with head; mesoscutum exposed; outer margins of five spicules, including spicules B and F (fig. 8J). hemelytra only slightly convex medially. Hem- Redescription: Male. Total length 3.9. elytron: Vein along behind clavus almost flat; COLORATION (fig. 5): Head: Uniformly yellow. inner margin of cuneus concave (as in Namya- Eye pale brown. Labium: Yellow. Thorax: Prono- tova et al., 2016: fig. 13E), outer margin of tum, mesoscutum, scutellum and thoracic cuneus ca. 3× as long as base. Abdomen: Genital pleura yellow; scent gland evaporative area whit- capsule rotated left at small angle relative to rest ish yellow, brown apically. Hemelytron: Mostly of abdomen. Genital capsule (fig. 14N) ca. 1.5× translucent, colorless; inner margin of clavus as long as wide; ventral wall ca. 1.5× as long as whitish yellow, with pale brown margin; area dorsal wall, its posterior margin smooth, semi- along inner margin of corium mostly yellow, oval, without outgrowth, not curved dorsally; its pale brown anteriorly; embolium and cuneus apex inclined rightward; sides of genital capsule with yellow tinge and with pale brown margins; not modified; right paramere socket slightly membrane cell yellow. Legs: Coxae, femora, and acute and left paramere rounded; distance tibiae yellow. Abdomen: Uniformly yellow. SUR- between paramere sockets subequal to half of FACE AND VESTITURE: Corium smooth, with genital capsule width at base. Right paramere scarce punctures. Dorsum with setae subequal (fig. 11AF) distinctly curved apically; medial to or slightly longer than antennal segment II part twice as wide as basal part, bearing setae, diameter; setae on femora subequal to or shorter with outer margin slightly concave; and inner than antennal segment II diameter. STRUC- margin convex; outer angle distinct, widened; TURE AND MEASUREMENTS: Body ca. 4.0× inner angle rounded, without setae; basal part of as long as pronotum width. Head: Depression right paramere ca. 0.15–0.2× as long as rest of delimiting occipital region present dorsally and paramere. Left paramere (fig. 11AG) L-shaped, laterally (as in Namyatova et al., 2016: fig. 4E), apical part flattened, with toothlike outgrowth distance between depression and pronotum dis- on posterior side medially (as in fig. 11G) and tinctly shorter than eye diameter; longitudinal without outgrowth on dorsal surface; middle sulcus on dorsal surface longer than eye diam- part widened, without swelling or outgrowth; eter; distance from eye to pronotum slightly lon- setae present only on middle part near outer ger than eye diameter, not swollen laterally (as margin. Aedeagus conjunctiva weakly sclero- in Namyatova et al. 2016: fig. 4E); vertex ca. 1.9× tized; secondary gonopore placed at base of as wide as eye; upraised (in Namyatova et al., vesica in repose; sclerotization of ductus seminis 2016: fig. 6D ); buccula ca. 0.2–0.25× as long as around secondary gonopore shorter than wide; clypeus. Labium: Surpassing middle of meso- vesica with five spicules, including spicules B sternum but not reaching posterior part of and F (fig. 8J). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 87
Female. Total length 4.0. COLORATION (fig. Material examined: Lectotype: SRI 5): Head: Yellow with reddish longitudinal red LANKA: Des Montagnes de Nuwara Eliya stripe behind eye and red stripe above antennal [Nura-Ellia] et Patannas., 6.9738°N 80.8265°E, fossae, tubercle around antennal fossa red later- 1700, Motschulsky, 1♂ (00017876) (NHRS). ally. Labium: As in male. Antenna: Segment I red Paralectotype: SRI LANKA: Des Montagnes de with yellow base, segment II reddish brown, seg- Nuwara Eliya [Nura-Ellia] et Patannas., 6.9738°N ment III brown. Eye brown. Thorax: Pronotum 80.8265°E, 1700, Motschulsky, paralectotype, 1♀ yellow with longitudinal red tripe laterally; tho- (00017871) (NHRS). racic pleura yellow; scent gland evaporative area whitish yellow basally and pale brown apically. Felisacus gressitti Miyamoto Hemelytron: Similar to male, but inner part of Figures 5, 18 clavus, embolium, and cuneus with red margins, Felisacus gressitti Miyamoto, 1965: 162 (original cuneus with reddish tinge, membrane cell red. description). Legs: Yellow, femora apically and entire tibiae with red stripe, hind tarsus with yellow segment Diagnosis: Recognized by the following com- I, segments II–III pale brown. SURFACE AND bination of characters: head and anterior part of VESTITURE: As in male; setae on antennal seg- pronotum mostly yellow, clypeus mostly dark ment I shorter than antennal segment II diame- brown; posterior part of pronotum dark brown ter. STRUCTURE AND MEASUREMENTS: with yellow stripe medially; antennal segment I Structure as in male; body ca. 3.9× as long as yellow, reddish apically; marking along inner mar- pronotum width; vertex ca. 2.0× as wide as eye; gin of corium brown, narrow, not extending antennal segment I cylindrical (as in Namyatova toward R+M anteriorly and posteriorly (fig. 5); et al., 2016: fig. 8A), ca. 1.3× as long as head antennal segment I slightly widened (as in Namya width, ca. 0.8× as long as pronotum width; seg- tova et al., 2016: fig. 6E); labium reaching abdomi- ment II ca. 1.8× as long as head width, ca. 1.0× nal segment II; antennal segment II ca. 1.6× as as long as pronotum width; pronotum ca. 1.3× as long as head width; right paramere sickle shaped wide as long and ca. 1.7× as wide as head. (Miyamoto, 1965: figs. 35, 36; Lin, 2000: fig. 2B) Distribution: Sri Lanka (fig. 18). and left paramere widened with outgrowth poste- Host plants: Unknown. riorly (Lin, 2000: fig. 2B). Discussion: We examined the male lectotype Redescription: Male. Not examined, see and female paralectotype of F. glabratus. The Miyamoto (1965) and Lin (2000). Female. Total specimens were damaged, and no information length 3.2. COLORATION (fig. 5): Mostly yel- has been recorded for the antennae and tarsi of low with reddish tinge posteriorly; tubercle male I, the antennal segment IV in female, col- around antennal fossa whitish yellow; clypeus oration of female abdomen, the apex of the right dark brown, red basally. Eye dark brown, red- paramere (broken). In addition, the female geni- dish laterally. Labium: Segments I–II whitish talia were not dissected. See generic discussion yellow, segments III–IV yellow. Antenna: Seg- for the explanation of the nomenclatural confu- ment I yellow with reddish tinge; segment II sion in literature. reddish brown; segments III–IV dark brown. Felisacus glabratus is closely related to F. albus, Thorax: Anterior part of pronotum yellow; pos- F. indicus, and F. insularis. They share a similar terior part of pronotum dark brown with yellow structure of the right paramere, which is broad stripe medially; scutellum, mesoscutum, and medially, twice as long as the basal part (figs. thoracic pleura yellow; scent gland evaporative 11A, AF, 12A, C). All of them also have similar area yellow basally and red apically. Hemelytron: vesical spicules, but differ in their shape and/or Mainly translucent and colorless; inner part of number (cf. fig. 8J with fig. 8A, K and L). corium opaque, yellow, brown basally and along 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 inner margin; area along inner margin of 3× as long as base. Female genitalia were not corium brown; embolium with yellow margins, investigated. brown apically; cuneus yellow; membrane with Distribution: South Japan (fig. 18). brown tinge; membrane cell brown. Legs: Coxae Host plants: Collected from ferns (Miya- whitish yellow; femora with whitish yellow moto, 1965). basal half and yellow apical half; tibiae yellow; Discussion: Characters of the male as tarsal segment I whitish yellow, segments II–III given in the diagnosis and discussion were pale brown to brown. Abdomen: Yellow, seg- adapted from Miyamoto (1965) and Lin ment IX reddish yellow with red lateral stripe. (2000). Miyamoto (1965) described Felisacus SURFACE AND VESTITURE: Corium with gressitti from two males and two females. shallow punctures. Dorsum and femora with According to his description, the holotype and setae longer than antennal segment II diameter; two paratypes are preserved in the Entomo- segment II clothed with suberect setae mostly logical Laboratory, Kyushu University, and a shorter than antennal segment II diameter; single paratype is preserved in BPBM. We did abdomen with setae of different length, mostly not locate the paratype in BPBM and we were longer than antennal segment II diameter. unsuccessful in acquiring information from STRUCTURE AND MEASUREMENTS: Body the Kyushu University on the types. ca. 4.1× as long as pronotum width. Head: We examined a single female from the Depression delimiting occipital region very Tomohide Yasunaga collection and it fits the shallow, present only dorsally; distance between description of F. gressitti. We did not dissect its depression and pronotum distinctly shorter genitalia: Felisacus gressitti is similar to F. viti- than eye diameter; longitudinal sulcus on dorsal levu externally (fig. 7) and they have a very surface of head slightly shorter than eye diam- similar paramere structure (cf. Miyamoto, 1965: eter; distance from eye to pronotum shorter figs. 34–38, Lin, 2000: fig. 2B; fig. 13V, W). than eye diameter, not swollen laterally; vertex However, F. vitilevu differs by the clypeus being ca. 1.7× as wide as eye; vertex upraised; buccula mostly yellow to pale brown, but is also some- ca. 0.2–0.25× as long as clypeus. Labium: Reach- times red, antennal segment II in males is ca. ing abdominal segment II; segments I and II 1.9–2.1× long as wide, and in female, it is ca. reduced, combined longer than half of segment 1.9–2.0× as long as wide. Although these two III; segment I shorter than wide; segment II species are consistently different, differences are longer than wide, its dorsal surface elongate minor their status is still in question, as we only posteriorly (as in Namyatova et al., 2016: fig. examined a single female of F. gressitti. None- 6E); segment IV twice as long as segment III. theless, they are broadly disjunct (F. gressitti is Antenna: Segment I slightly widened medially, known from Japan and Taiwan and F. vitilevu ca. 1.3× as long as head width, ca. 0.8× as long inhabits Fiji Islands), and pending further evi- as pronotum width; segment II ca. 1.6× as long dence, we treat them as distinct. as head width, ca. 1.0× as long as pronotum Felisacus gressitti is also similar to F. mada- width; segment III subequal to segment II. Tho- gascariensis in structure and coloration (fig. rax: Anterior part of pronotum subequal to half 5), but the latter species differs by the prono- of posterior part; posterior part slightly tum being brown posteriorly, the marking upraised; posterior margin straight; pronotum along the inner margin of corium is yellow, ca. 1.3× as wide as long and ca. 1.6× as wide as and the labium reaches the posterior margin of head; mesoscutum exposed. Hemelytron: Area the mesosternum. along inner margin of corium almost flat; inner Material examined: JAPAN: Okinawa: margin of cuneus convex (as in Namyatova et Ohshittai, Nago, 26.59336°N 127.97844°E, 02 al., 2016: fig. 13E), outer margin of cuneus ca. Dec 2001, T. Yasunaga, 1♀ (00201839) (TYCN). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 89
Felisacus indicus, sp. nov. apically; hind tibia whitish with reddish tinge; tarsi whitish, pale brown dorsally. Abdomen: Figures 5, 8K, 12A, B, 14O, 18 Uniformly whitish yellow. SURFACE AND Diagnosis: Recognized by the following set VESTITURE: Corium smooth, often with scarce of characters: coloration of dorsum mostly whit- shallow punctures. Dorsum with suberect setae ish yellow, pronotum with brown humeral angles; most often slightly longer than diameter of cuneus opaque, whitish yellow with yellow apex antennal segment II; antennal segment I and and margins (fig. 5), cylindrical antennal seg- femora with suberect setae shorter than diam- ment I (as in Namyatova et al., 2016: fig. 8A), eter of antennal segment II; abdomen clothed transverse depression on head delimiting occipi- with suberect short simple setae. STRUCTURE tal region extending laterally, vertex in males ca. AND MEASUREMENTS: Body ca. 4.3× as long 1.2× as wide as eye, upraised (as in Namyatova et as pronotum width. Head: Depression delimit- al., 2016: fig. 6D); labium reaching middle of ing occipital region present dorsally and later- metasternum; cuneus ca. 2–2.5× as long as its ally (as in Namyatova et al., 2016: fig. 4E); width at base; genital capsule almost as long as distance between depression and pronotum dis- wide (fig. 14O); medial part of right paramere tinctly shorter than eye diameter; longitudinal twice as wide as basal part, and shorter than api- sulcus on dorsal surface longer than eye diam- cal and basal parts combined; its outer margin eter; distance from eye to pronotum slightly straight and inner margin widened (fig. 12A); longer than eye diameter, not swollen laterally vesica with five spicules, including spicules B and (as in Namyatova et al., 2016: fig. 4E); vertex ca. F, without serrate spicules (fig. 8K). 1.2× as wide as eye; vertex upraised; buccula ca. Description: Male. Total length 4.0. COL- 0.2–0.25× as long as clypeus. Labium: Reaching ORATION (fig. 5): Head: Mostly whitish yellow middle of metasternum; segments I and II with reddish markings. Eye whitish yellow. strongly reduced, combined length shorter than Labium: Segments I–III whitish yellow, segment segment III; segment I slightly shorter than IV yellow. Antenna: Segments I–II whitish yel- wide (as in Namyatova et al., 2016: figs. 6D, 9C), low with reddish tinge, segments III–IV pale segment II slightly longer than wide, elongate brown. Thorax: Pronotum whitish yellow with dorsally, segment IV ca. 1.5× as long as segment brown anterior margin and posterior angles; III. Antenna: Segment I cylindrical (as in Namy- punctures between anterior and posterior parts atova et al., 2016: fig. 8A), ca. 1.3× as long as pale brown; mesoscutum and scutellum whitish head width, ca. 0.9× as long as pronotum width; yellow with pale brown punctures between segment II ca. 1.7× as long as head width, ca. them; thoracic pleura whitish yellow; scent 1.1× as long as pronotum width. Thorax: Ante- gland evaporative area whitish basally and pale rior part of pronotum only slightly shorter than brown apically. Hemelytron: Inner part of clavus posterior part; collar delimited; posterior part opaque, whitish yellow, with brown margins; upraised; posterior margin concave; pronotum outer part of clavus whitish and translucent; ca. 1.2× as wide as long and ca. 1.5× as wide as corium translucent, area along inner margin of head; mesoscutum exposed. Hemelytron: Area corium pale brown anteriorly and whitish yel- along inner margin on corium almost flat; inner low posteriorly; embolium opaque, whitish yel- margin of cuneus convex, costal margin ca. low, with apex and margins pale brown; cuneus 2–2.5× as long as base. Abdomen: Genital cap- opaque, whitish yellow, with yellow apex and sule weakly rotated to left-hand side. Genitalia: margins; membrane translucent, colorless; Genital capsule (fig. 14O) almost as long as membrane cell mostly whitish yellow, pale wide; ventral wall twice as long as dorsal wall, brown anteriorly. Legs: Coxae whitish; femora its posterior margin smooth, semioval, without and tibiae mostly whitish yellow, hind femur red outgrowth, not curved; left side of genital cap- 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 sule not widened; right paramere socket slightly Felisacus insularis Miyamoto acute and left paramere socket rounded; dis- Figures 5, 8L, 12C, D, 14P, 18 tance between paramere sockets subequal to half of genital capsule with at base. Right para- Felisacus insularis Miyamoto, 1965: 159 (original mere (fig. 12A) distinctly curved in apical half; description). apex almost straight posteriorly; medial part Diagnosis: Recognized by the following twice as wide as basal part, bearing setae, with combination of characters: mostly yellow color- outer margin straight and inner margin convex; ation of head and pronotum, head, collar, and outer angle distinct; inner angle rounded, with- inner part of clavus without brown markings; out setae; basal part of right paramere ca. 0.15– cuneus often at least partly yellow or red (fig. 5), 0.2× as long as rest of paramere; apical part of cylindrical antennal segment I (as in Namyatova right paramere lost. Left paramere (fig. 12B) et al., 2016: fig. 8A), transverse depression delim- L-shaped; apical part not flattened, with tooth- iting occipital region on head extending laterally, like outgrowth on posterior side medially (as in vertex upraised (as in Namyatova et al., 2016: fig. fig. 11G) and without outgrowth on dorsal sur- 6D); body length in male 3.7–4.2 and in female face; middle part widened, without swelling or 3.8–4.3; vertex in male ca. 1.5–1.9× as long as eye outgrowth; setae only on middle part near outer diameter; labium reaching middle of mesoster- margin. Aedeagus conjunctiva weakly sclero- num; ventral wall of genital capsule ca. 1.7× as tized; secondary gonopore placed at base of long as dorsal wall (fig. 14P); medial part of right vesica in repose; sclerotization of ductus semi- paramere twice as wide as basal part, shorter nis around secondary gonopore of shorter than than basal and apical parts combined, its outer wide; vesica with five spicules including spicules B and F (fig. 8K). margin straight (fig. 12C); vesica with five spic- Female. Unknown. ules, including spicules B and F and large serrate Distribution: India (fig. 18). spicule (fig. 8K). Host plants: Unknown. Redescription: Male. Total length 3.7–4.2. Discussion: Felisacus indicus is similar to F. COLORATION (fig. 5): Head: Mostly whitish albus in external morphology (figs. 4, 5), and yellow to yellow; frons, mandibular plate and the structure of the genital capsule (fig. 14A, O) tubercle around antennal fossa often with red- and parameres (cf. fig. 11A, B with fig. 12A, B). dish tinge or small markings. Eye dark brown, Felisacus albus differs from F. indicus by the often reddish at margins. Labium: Yellow, seg- humeral angles of the pronotum whitish yellow, ments I–II sometimes whitish yellow, segment II the cuneus is whitish yellow to yellow with a with pale brown or reddish stripe ventrally. reddish tinge and the presence of seven vesical Antenna: Segment I yellow to pale brown, whit- spicules (fig. 8A). Felisacus indicus is also simi- ish yellow basally, sometimes with reddish tinge; lar to F. insularis in color patterning (fig. 5), the segment II reddish brown, darker than segment shape of the right paramere (cf. fig. 21A with I, segment III–IV brown to dark brown, darker 12C) and the presence of spicules B and F (cf. than segment II. Thorax: Pronotum whitish yel- fig. 8K with 8L). The latter species differs by the low to yellow, sometimes with reddish stripe lat- head and the humeral angles yellow, the ventral erally, anterior margin often pale brown, wall of the genital capsule ca. 1.7× as long as the sometimes humeral angle pale brown; thoracic dorsal wall (fig. 14P), and the vesica has a ser- pleura whitish yellow to yellow, scent gland rate spicule (fig. 8L). evaporative area whitish yellow often yellow api- Material Examined: Holotype: INDIA: cally, sometimes yellow with pale brown apex; Kerala: Kerala Prov, 10°N 76.5°E, 1991, P.K. mesoscutum and scutellum whitish yellow to Sumodan, 1♂ (00017872) (NML). yellow, punctures between them often pale 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 91 brown. Hemelytron: Mostly translucent, color- long as pronotum width; segment II ca. 1.5–2.2× less; inner part of clavus whitish yellow to yellow, as long as head width, ca. 0.9–1.4× as long as with yellow or pale brown margins, sometimes pronotum width; segments III slightly longer opaque; corium with marking along inner mar- than segment II; segment IV ca. 0.3× as long as gin yellow to pale brown, narrow and short; segment III. Thorax: Posterior part slightly embolium sometimes yellow, with brown mar- upraised; collar delimited posterior margin gins, often reddish or reddish brown apically; slightly concave; pronotum ca. 1.2–1.4× as wide cuneus yellow to reddish yellow, sometimes only as long and ca. 1.5–1.7× as wide as head; meso- apex yellow or reddish or cuneus with yellow scutum exposed. Hemelytron: Area along inner tinge only; membrane cell yellow. Legs: Coxae margin of corium flat; inner margin of cuneus whitish yellow to yellow; femora whitish yellow convex, outer margin of cuneus ca. 2.5–3× as basally and yellow apically, rarely mostly pale long as base. Abdomen: Genital capsule rotated brown, often with reddish tinge apically; tibiae left at right angle relative to rest of abdomen. yellow to pale brown, often reddish basally, rarely Genitalia: Genital capsule (fig. 14P) twice as long mostly reddish; tarsi yellow to pale brown. Abdo- as wide, ventral wall ca. 1.5× as long as dorsal men: Ventral and lateral sides yellow, segment IX wall, its posterior margin smooth, semioval, pale brown, dorsal surface reddish, rarely abdo- without outgrowth(s), not curved; sides of geni- men uniformly yellow. SURFACE AND VESTI- tal capsule not modified; paramere socket more TURE: Corium smooth, with shallow and scarce or less rounded; distance between paramere punctures. Dorsum with suberect setae slightly sockets ca. 0.7× as long as genital capsule width longer than antennal segment II diameter; anten- at base; right angle of left paramere socket not nal segment I and femora with suberect setae projecting. Right paramere (fig. 12C) distinctly subequal to or shorter than antennal segment II curved in apical half; apex straight posteriorly; diameter; abdomen clothed with suberect mostly medial part almost twice as wide as basal part, short simple setae. STRUCTURE AND MEA- bearing setae, with outer margin straight and SUREMENTS: Body ca. 3.8–4.3× as long as pro- inner margin widened; outer angle distinct, notum width. Head: Depression delimiting inner angle rounded, without setae; basal part of occipital region present dorsally and laterally (as paramere ca. 0.15–0.2× as long as rest of param- in Namyatova et al., 2016: fig. 4E); distance ere. Left paramere (fig. 12D) L-shaped; apical between depression and pronotum distinctly part not flattened, with toothlike outgrowth on shorter than eye diameter; longitudinal sulcus on posterior side medially (as in fig. 11G) and with- dorsal surface longer than eye diameter; distance out outgrowth on dorsal surface; middle part from eye to pronotum slightly longer than eye widened, without swelling or outgrowth; setae diameter, not swollen at sides (Namyatova et al., only on middle part near outer margin. Aedeagus 2016: fig. 4E); vertex ca. 1.5–1.9× as wide as eye, conjunctiva weakly sclerotized; secondary gono- upraised (as in Namyatova et al., 2016: fig. 6D). pore placed at base of vesica in repose; sclerotiza- Labium: Reaching middle of mesosternum; seg- tion of ductus seminis around secondary ments I and II strongly reduced, combined sub- gonopore shorter than wide; vesica with five equal to half of segment III; segment I slightly spicules, including spicules B and F and large shorter than wide (as in Namyatova et al., 2016: serrate spicule (fig. 8L). figs. 6D, 9C); segment II slightly longer than Female. Total length 3.8–4.3. COLORATION wide, elongate dorsally; segment III as long as (fig. 5): Similar to male, abdomen: uniformly yel- ventral side of head length; segment IV ca. 1.5× low, sometimes with dorsal surface pale brown. as long as segment III. Antenna: Segment I cylin- SURFACE AND VESTITURE: As in male. drical (as in Namyatova et al., 2016: fig. 8A), ca. STRUCTURE AND MEASUREMENTS: Struc- 1.2–1.8× as long as head width, ca. 0.8–1.1× as ture as in male; body ca. 4.0–4.6× as long as pro- 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 notum width; vertex ca. 1.5–1.9× as wide as eye; seven vesical spicules (fig. 8A). Felisacus indicus antennal segment I ca. 1.3–1.4× as long as head can be separated by the vertex in males being ca. width, ca. 0.7–0.9× as long as pronotum width; 1.2× as wide as the eye diameter, the genital cap- segment II ca. 1.7–1.9× as long as head width, ca. sule is almost as long as wide (fig. 14O), the 1.0–1.2× as long as pronotum width; pronotum humeral angles of the pronotum are brown, and ca. 1.1–1.5× as wide as long and ca. 1.5–1.8× as the vesica is without serrate spicules (fig. 8L). wide as head. Genitalia (as in Namyatova et al., Material examined: CHINA: Guangdong: 2016: fig. 23F, G): Dorsal labiate plate wider than Yao’anxiang, Lianzhou, Qingyuan, 25.01826°N distance between apodemes of second valvula; 112.40858°E, 20 Oct 1962, Hanhua Cheng and mostly smooth, without distinct striations or lon- Leyi Zheng, 1♂ (00018484) (NKMU). Hainan gitudinal membranous ridge, with semicircular Prov.: Jianfeng Mt, 18.72°N 108.88°E, 08 May sclerites and distinct sclerotized rings laterally; 1964, Unknown, 1♂ (00018482) (NKMU); Apr lateral oviducts placed almost medially, very close 1980, Unknown, 1♂ (00017887), 2♀ (00018480, to each other, spermathecal gland placed between 00018481) (NKMU). Yunnan: [Man Xue], 06 lateral oviducts; dorsal labiate plate with distinct Jun 1958, Unknown, 1♀ (00018483) (IOZ). tubercles, without membranous lobe medially. JAPAN: Kyushu (Island): Okinawa Pref.: Ish- Distribution: China, Japan, Thailand, Viet- igaki I., Kara-yama, 14 Mar 1964, C.M. Yoshi- nam (fig. 18). moto and J. Harrell, 2♀ (00018486, 00018479) Host plants: Sweeping from Pteridium sp. (BPBM). Okinawa: Ishigaki Is., Ryukyu Islands, (Miyamoto, 1965). 24.4°N 124.2°E, Oct 1951, R.M. Bohart, 1♂ Discussion: Miyamoto (1965) described F. (00018477) (AM); 13 Oct 1999, Belokobylskij, insularis based on the basis of a male holotype 3♀ (00271508, 00271507), 2♂ (00271509, and a number of male and female paratypes. 00271508) (ZISP). THAILAND: Chiang Mai According to information given in his descrip- Province: Doi Suthep, 18.81277°N 98.89361°E, tion, the holotype and some paratypes are pre- 900 m, 14 Nov 1957, J.L. Gressitt, 1♂ (00017875) served in the Entomological Laboratory, Kyushu (AM). VIETNAM: Gia Lai: 20 km N of Pleiku, University, and two paratypes are preserved in 14.15162°N 107.94731°E, 650 m, 09 May 1960, the BPBM. We did not find the paratypes of F. L.W. Quate, 1 sex unknown (00017866), 1♂ insularis in BPBM, and we did not acquire infor- (00017874) (AM). Ha Noi: 70 km NWW Ha Noi mation about the types of this species from BaVi, 21.087°N 105.302°E, 22 Nov 1990, Beloko- Kyushu University. bylskij, 1♂ (00018478) (AM). We have examined a number of specimens from southeast Asia belonging to a single species Felisacus jacobsoni Poppius and more or less fitting the description of Miya- Figures 5, 8M, 12E, F, 18 moto, aside from minor variation in coloration. Felisacus jacobsoni Poppius, 1914: 149 (original These specimens also have a similar right para- description). mere, with a broad medial region, as depicted in Miyamoto (1965: fig. 30), and we consider them Diagnosis: Recognized by the following as conspecific to F. insularis. combination of characters: dorsal surface of head Felisacus insularis is similar to F. albus and F. brown anteriorly; pronotum brown along poste- indicus (figs. 4, 5) in external view and the medial rior margin; inner part of clavus opaque, brown; part of the right paramere, which is twice as wide scent gland evaporative area brown; cuneus red as the basal part (cf. fig. 12C with figs. 11A, 12A). with outer margin brown; marking along inner Felisacus albus differs by the mostly whitish yel- margin of corium brown, not reaching R+M low head and collar, the genital capsule is almost anteriorly and posteriorly (fig. 5), cylindrical as long as wide (fig. 14A), and the presence of antennal segment I (as in Namyatova et al., 2016: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 93 fig. 8A), transverse depression on head extending length subequal to half of segment III; segment laterally, vertex upraised (as in Namyatova et al., I shorter than wide; segment II slightly longer 2016: fig. 6D); labium reaching posterior margin than wide, its dorsal surface elongate posteri- of metasternum; vesica with five spicules, includ- orly; segment III slightly shorter than ventral ing spicules A and B, spicule A elongate and wid- side of head; segment IV twice as long as seg- ened, spicule B small and not swollen (fig. 8M). ment III. Antenna: Segment I cylindrical (as in Redescription: Male. Total length 4.4–4.7. Namyatova et al., 2016: fig. 8A), ca. 1.6× as long COLORATION (fig. 5): Head: Mostly yellow, as head width, ca. 1.0–1.1× as long as pronotum brown dorsally, labrum brown. Eye dark brown. width; segment II ca. 2.1–2.3× as long as head Labium: Uniformly yellow. Antenna: Segment I width, ca. 1.4× as long as pronotum width; seg- brown with yellow base, segments II–IV dark ments III slightly longer than segment II; seg- brown. Thorax: Pronotum yellow with brown ment IV ca. 0.3× as long as segment IV. Thorax: collar and markings on anterior part of prono- Anterior part of pronotum slightly shorter than tum laterally, posterior part of pronotum with posterior part; posterior part of pronotum paired brown markings near margin; scutellum slightly upraised; posterior margin of pronotum yellow; pleura yellow, scent gland evaporative concave, pronotum ca. 1.2–1.3× as wide as long area brown, with whitish yellow apex and base. and ca. 1.6× as wide as head; mesoscutum Hemelytron: Mostly translucent and colorless; exposed. Hemelytron: Area behind clavus almost inner part of clavus opaque, brown; marking flat; inner margin of cuneus convex (as in along inner margin of corium brown, cuneus Namyatova et al., 2016: fig. 13E), outer margin mostly red with outer margin translucent with of cuneus almost ca. 3× as long as base. Genita- yellow tinge and margins pale brown; mem- lia: Right paramere (fig. 12E) apical part dis- brane with grayish tinge and pale brown veins. tinct; apex slightly concave; medial part less Legs: Coxae whitish yellow; femora whitish yel- than twice as wide as basal part, bearing setae, low basally and yellow apically; tibiae yellow to with outer margin straight and inner margin pale brown, brown basally; tarsi brown. SUR- convex; outer angle distinct, not widened; inner FACE AND VESTITURE: Corium smooth with angle rounded, without setae; basal part ca. 0.2– shallow and scarce punctures. Dorsum and 0.25× as long as rest of paramere. Left paramere femora with suberect setae mostly subequal to (fig. 12F) L-shaped; apical part not flattened, antennal segment II diameter; antennal segment with toothlike outgrowth on posterior side I with scarce suberect setae mostly shorter than medially (as in fig. 11G) and without outgrowth antennal segment II diameter. STRUCTURE on dorsal surface; middle part widened, without AND MEASUREMENTS: Body ca. 4.4–4.6× as swelling or outgrowth(s); setae only on middle long as pronotum width. Head: Depression, part near outer margin. Aedeagus conjunctiva delimiting occipital region, present dorsally and weakly sclerotized; secondary gonopore placed laterally (as in Namyatova et al., 2016: fig. 4E); at base of vesica in repose; sclerotization of duc- distance between depression and pronotum dis- tus seminis around secondary gonopore shorter tinctly shorter than eye diameter; longitudinal than wide; vesica with five spicules: spicule A is sulcus on dorsal surface longer than eye diam- long and wide, spicule B short; three additional eter; distance from eye to pronotum slightly lon- spicules present on left-hand side, all narrow ger than eye diameter, not swollen laterally (as and acute (fig. 8M). in Namyatova et al., 2016: fig. 4E); vertex ca. Female. Total length 4.6–4.8. COLORATION 1.4–1.7× as wide as eye upraised. Labium (as in (fig. 5): Similar to male, head often dark brown Namyatova et al., 2016: figs. 6D, 9C): Almost dorsally, sometimes with reddish-brown mark- reaching posterior margin of metasternum; seg- ing between eyes; pronotum yellow sometimes ments I and II strongly shortened, combined with brown stripe along posterior margin; tho- 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 racic pleura sometimes with brown markings or Material examined: Lectotype: INDONE- mostly pale brown; tibiae sometimes uniformly SIA: East Java: Nongkojajar [Nongkodjadjar], yellow to pale brown; tarsal segments I–II some- 7.8965°S 112.8213°E, 1095 m, Jan 1911, E. Jacob- times whitish yellow. SURFACE AND VESTI- son, 1♀ (00017877) (MZH). Additional material: TURE: As in male. STRUCTURE AND West Nusa Tenggara: Lombok Island, nr Senaru, MEASUREMENTS: Similar to male. body ca. 8.31956°S 116.405°E, 01 Sep 2012, F. Konstanti- 4.3–5.1× as long as pronotum width; vertex ca. nov, Pteris sp. (Pteridaceae), det. Michael Lovave 1.7–1.8× as wide as eye; antennal segment I ca. (LAE herbarium, PNG), 2♂ (00386513, 1.5–1.7× as long as head width, ca. 0.9–1.1× as 00386514), 4♀ (00386515–00386518) (ZISP). long as pronotum width; segment II ca. 2.1–2.2× as long as head width, ca. 1.3–1.4× as long as Felisacus javanus (Reuter) pronotum width; pronotum ca. 1.1–1.2× as wide Figures 5, 8N, 12G, H, 14Q, 19 as long and ca. 1.4–1.6× as wide as head. Genita- Hyaloscytus elegantulus var. javanus Reuter, lia (as in fig. 23F, G): Dorsal labiate plate wider 1908: 187 (original description), stat. nov. than distance between apodemes of second val- Felisacus carpenterae Hsiao, 1944: 385, new vula; mostly smooth, without distinct striations, synonymy. with semicircular sclerite and distinct sclerotized rings laterally; lateral oviducts placed almost Diagnosis: Recognized by the following medially, very close to each other, spermathecal combination of characters: main coloration of gland placed between lateral oviducts; dorsal head and pronotum yellow to pale brown, with- labiate plate with distinct tubercles, without out brown markings, cuneus with inner part red membranous lobe medially. and outer part whitish yellow, margins red (fig. Distribution: Java and Lombok Is. (Indone- 5), cylindrical antennal segment I (as in Namya- sia) (fig. 18). tova et al., 2016: fig. 8A), transverse depression Host plants: Pteris sp. on head delimiting occipital region extending Discussion: Poppius (1914) described Feli- laterally, vertex upraised (as in Namyatova et al., sacus jacobsoni from a single female, collected in 2016: fig. 6D) body length in male 3–3.7, in Java and preserved in the FMNH. Some speci- female 3.5–4.1; labium reaching posterior mar- mens in the collection of ZISP are from Lombok gin of mesosternum or slightly surpassing it; are very similar to the type specimen, and we cuneus ca. 2.5× as long as base; ventral wall of assigned them to this species. genital capsule ca. 1.5× as long as dorsal wall (fig. Felisacus jacobsoni is most similar to F. bellus, 14Q); medial part of right paramere slightly F. bau, and F. signis in coloration (fig. 4): Feli- wider than basal part, shorter than basal and api- sacus bau and F. signis differ in body size, which cal parts combined, its outer margin concave is less than 4 mm in both sexes. All these species (fig. 12G); vesica with five spicules, including differ in the shape of the vesical spicules (cf. fig. spicules A, B, and C; spicule A longer than spic- 8M with figs. 8D, E, 9K). ule B, spicule B long and swollen, spicule C long Felisacus jacobsoni can be confused with F. and distinctly convolute (fig. 8N). javanus (fig. 5), which is widely distributed in Redescription: Male. Total length 3.0–3.7. Indonesia, as the latter also has a bright red COLORATION (fig. 5): Head: Yellow to pale cuneus and brown markings. However, F. java- brown, dorsal surface and clypeus sometimes nus is paler and its head and pronotum have with reddish tinge. Eye dark brown to black, brown markings, it is smaller, body length, which sometimes with pale brown spots or reddish is 3.0–3.7 mm in males and 3.5–4.1 mm in tinge. Labium: Whitish yellow to pale brown, females, and has distinct vesical spicules with segment III sometimes with reddish tinge ven- spicule B elongate and broad (fig. 8N). trally. Antenna: Segment I pale brown, some- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 95 times with reddish tinge, segment II reddish et al., 2016: fig. 4E); vertex ca. 1.3–1.9× as wide brown, segments III dark brown. Thorax: Prono- as eye, upraised (as in Namyatova et al., 2016: fig. tum yellow with anterior margin brown, some- 6D). Labium (as in Namyatova et al., 2016: figs. times whitish yellow posteriorly, sometimes with 6D, 9C): Reaching posterior margin of mesoster- reddish tinge laterally; humeral angle or marking num or slightly surpassing it; segments I and II nearby often yellow to pale brown; mesoscutum strongly reduced, combined subequal to half of and scutellum whitish yellow to pale brown, segment III; segment I shorter than wide; seg- punctures between them pale brown, scutellum ment II slightly longer than wide, its dorsal sur- with longitudinal brownish stripe; thoracic face elongate posteriorly; segment III as long as pleura yellow to pale brown, sometimes with ventral side of head; segment IV ca. 1.5× as long reddish tinge; scent gland evaporative area yel- as segment III. Antenna: Segment I cylindrical low, sometimes red apically. Hemelytron: Mostly (as in Namyatova et al., 2016: fig. 8A), ca. 1.4– translucent; clavus sometimes opaque, inner part 1.6× as long as head width, ca. 1.0–1.2× as long of clavus whitish yellow to yellow, sometimes as pronotum width; segment III slightly longer brown apically, with yellow, reddish or pale than segment II. Thorax: Anterior part of prono- brown margins; outer part of clavus colorless or tum only slightly shorter than posterior part; col- whitish yellow, with apex yellow to brown; lar delimited; posterior part slightly upraised; corium whitish yellow to yellow, area along inner posterior margin concave, pronotum ca. 1.1– margin of corium yellow to pale brown at middle 1.2× as wide as long and ca. 1.4–1.5× as wide as part and reddish posteriorly; embolium yellow head; mesoscutum exposed. Hemelytron: Vein with pale brown margins and often with reddish along behind clavus almost flat; inner margin of margin; cuneus with inner part red and outer cuneus convex (as in Namyatova et al., 2016: fig. part whitish yellow, margins and often its inner 13E), outer margin of cuneus ca. 2.5× as long as part red; membrane with yellow or grayish tinge, base. Genitalia: Genital capsule (fig. 14Q) ventral cell red. Legs: Coxa whitish yellow to yellow; wall ca. 1.5× as long as dorsal wall, with poste- femora whitish yellow to yellow basally and yel- rior margin of ventral wall semioval, smooth, low to pale brown apically, sometimes with red- without outgrowth(s), its apex inclined leftward, dish tinge; tibia yellow to pale brown, sometimes not curved; sides of genital capsule not modified; somewhat paler basally, sometimes with reddish right paramere socket slightly acute, left para tinge basally; tarsi pale brown, segment I often mere socket rounded; distance between param- whitish yellow to yellow. SURFACE AND VES- ere sockets subequal to half of genital capsule TITURE: Corium smooth, with shallow and width at base. Right paramere (fig. 12G) dis- scarce punctures. Dorsum with setae shorter or tinctly curved in apical half; apex almost straight; subequal to antennal segment II diameter; anten- medial part only slighter wider than basal part, nal segment I and femora clothed with suberect with row of setae, with outer margin slightly con- setae mostly shorter than width of antennal seg- cave and inner margin widened; outer angle dis- ment II diameter. STRUCTURE AND MEA- tinct, slightly swollen; inner angle rounded, not SUREMENTS: Body ca. 4.0–4.4× as long as bearing setae; basal part of paramere ca. 0.15– pronotum width. Head: Depression delimiting 0.2× as long as rest of paramere. Left paramere occipital region present dorsally and laterally (as (fig. 12H) L-shaped; apical part flattened, with in Namyatova et al., 2016: fig. 4E); distance tooth on posterior side medially (as in fig. 11G) between depression and pronotum distinctly and without outgrowth on dorsal surface; middle shorter than eye diameter; longitudinal sulcus on part widened, without swelling or outgrowth; dorsal surface of head longer than eye diameter; setae only on middle part near outer margin. distance from eye to pronotum longer than eye Aedeagus (general view as in Namyatova et al., diameter, not swollen laterally (as in Namyatova 2016: fig. 22I) conjunctiva weakly sclerotized, 96 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
A
F. javanus F. lindbergae F. lordhowensis F. luzonus
B
F. liui F. malayensis F. meilingae
C
F. longiceps F. myersi F. ochraceus F. ovalau
FIGURE 19. Distribution of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 97 secondary gonopore placed at base of vesica in sacus carpenterae was described from Singapore repose; sclerotization of ductus seminis around (Hsiao, 1944) and its type specimens are pre- secondary gonopore shorter than wide; vesica served in the USNM, and we examined male and with five long spicules, vesica with five spicules, female paratypes. On the basis of very similar including spicules A, B, and C; spicule A long external morphology and identical male genita- and wide, spicule B long and swollen, spicule C lia, we synonymize F. carpenterae with F. javanus. long and distinctly convolute (fig. 8N). In contrast, F. carpenterae signis Hsiao, 1944, dif- Female. Total length 3.5–4.1. COLORATION fers from the nominotypical subspecies and rep- (fig. 5): Head: Similar to male, but sometimes with resents a separate species (see the discussion of stripe above antennal fossae, markings between F. signis). See generic discussion for the explana- antennal fossae and stripe or marking behind eye tion of the nomenclatural confusion concerning red. Labium and antenna: As in male. Thorax: F. javanus in literature. Similar to male, sometimes pale brown with Felisacus javanus is most similar to F. amboi- brown anterior margin, scent gland evaporative nae, F. filicicola, F. ochraceus, and F. solomonicus area sometimes yellow with pale brown margin. in coloration and structure. Felisacus amboinae, Hemelytron: Similar to male, embolium rarely pale F. filicicola, and F. solomonicus can be separated brown. Legs: Similar to male, rarely uniformly by the right paramere with the outer margin pale brown. Abdomen: Whitish yellow with red- straight (cf. fig. 12G with figs. 11C, AD, 13R) and dish dorsum. SURFACE AND VESTITURE: As in the shape of the vesical spicules (cf. fig. 8N with male. STRUCTURE AND MEASUREMENTS: figs. 8B, I, 9L). Felisacus ochraceus is similar to F. Structure as in male; body ca. 4.0–4.4× as long as javanus in the shape of the right paramere and width of pronotum; vertex ca. 1.3–2.0× as wide as the configuration of the spicules (figs. 9F, 13C). eye; antennal segment I ca. 1.3–1.5× as long as In contrast, F. ochraceus has spicule B longer head width, ca. 0.9–1.0× as long as pronotum than spicule A. width; antennal segment II ca. 1.7–1.9× as long as Felisacus javanus and F. lambkinae are near head width, ca. 1.1–1.3× as long as pronotum identical in the shape and configuration of the width; pronotum ca. 1.2–1.4× as wide as long and vesical spicules (cf. fig. 8N with 8O), body size ca. 1.5–1.6× as wide as head. Genitalia (as in and structure of the right paramere, with both Namyatova et al., 2016: fig. 23F, G): Dorsal labiate having the medial region elongate, and the outer plate wider than distance between apodemes of margin is concave (cf. fig. 12G with 12I).Felisacus second valvula; mostly smooth, without distinct lambkinae differs from F. javanus by the anterior striations, with semicircular sclerite and distinct part of the pronotum being brown, the inner part sclerotized rings laterally; lateral oviducts placed of the clavus is pale brown to brown (fig. 5), and almost medially, very close to each other, sperma- the labium reaches or slightly surpasses the poste- thecal gland placed between lateral oviducts; dor- rior margin of the metasternum. sal labiate plate with distinct tubercles, without Felisacus javanus can be confused with F. membranous lobe medially. jacobsoni as the two species can be collected Distribution: Indonesia (Java, West Papua), together and are similar to each other in color- Malaysian Borneo (Sabah, Sarawak), Singapore ation, including the pale brown ground color, (fig. 19). presence of brown markings, with the cuneus at Host plants: Nephrolepis biserrata least partly red. However, in contrast to F. java- (Davalliaceae). nus, specimens of F. jacobsoni have a longer Discussion: Felisacus javanus was described body, with males 4.4–4.7 mm and females 4.6– from a series of specimens from Java (Reuter, 4.8 mm in length and, the labium reaches the 1908), and are preserved in the MZH. We exam- posterior margin of the mesosternum. Felisacus ined the male and female of the type series. Feli- jacobsoni differs slightly in coloration, having the 98 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 inner part of the clavus, scent gland evaporative 110.74222°E, 04 Jul 1958–09 Jul 1958, T.C. Maa, area, and anterior part of the head brown and in 1♀ (00021535) (ANIC), 3♂ (00021522, vesical spicules (fig. 8M). 00021533, 00021534) (BPBM). Material examined: Lectotype (F. javanus): INDONESIA: Java: collector unknown, 1902, Felisacus lambkinae, sp. nov. O.M. Schmiedeknecht, Lectotype, 1♂ (00018430) (MZH). Paralectotypes (F. javanus). INDONE- Figures 5, 8O, 12I, J, 14R, 18 SIA: Java: collector unknown, 1902, O.M. Diagnosis: Recognized by anterior part of Schmiedeknecht, paralectotype, 1♂ (00018435), pronotum brown; inner part of clavus brown to 4♀ (00018431–00018434) (MZH). Paratypes (F. pale brown; marking along inner margin of carpenterae). SINGAPORE: Singapore: Singa- corium brown, not reaching R+M; cuneus red- pore, 1.291°N 103.848°E, no date provided, C.F. dish with colorless and translucent outer part Baker, 1♂ (00338826), 1♀ (00338825) (USNM). and brown outer margin; cylindrical antennal Additional material: INDONESIA: West Papua: segment I (as in Namyatova et al., 2016: fig. 8A), Biak Island, W of Airport, 1.3425°S 136.1875°E, transverse depression on head extending later- 12 Aug 2012, F. Konstantinov, 2♀ (00271384, ally, vertex upraised; small size, body length in 00271385) (ZISP). Biak Island nr Taman Burung, male 3.1–3.2 and in female 3.2–3.5; labium 1.17603°S 136.18756°E, 10 Aug 2012, F. Konstan- reaching posterior margin of metasternum or tinov, Nephrolepis biserrata (Sw.) Schott (Daval- slightly surpassing it; vertex in male ca. 1.2–1.3× liaceae), det. Michael Lovave (LAE herbarium, as long as eye diameter; cuneus ca. 2.5× as long PNG), 11♂ (00271364–00271374), 3 juvenile wide; medial part of right paramere almost as (00271375–00271377), 6♀ (00271378– wide as basal part, its length subequal to basal 00271383) (ZISP). Manokwari, nr Taman and apical parts combined; its outer margin con- Gunung, Meja, 0.84497°S 134.07342°E, 21 Aug cave and inner margin convex (fig. 12I); vesica 2012, F. Konstantinov, 5♂ (00271389–00271393), with five spicules, including spicules A, B, and C 3♀ (00271395–00271397), 1 sex unknown (fig. 8O). (00271394) (ZISP). Owi Island nr Biak Island, Description: Male. Total length 3.1–3.2. 1.23817°S 136.20294°E, 12 Aug 2012, F. Konstan- COLORATION (fig. 5): Head: Dorsal surface tinov, Nephrolepis biserrata (Sw.) Schott (Daval- brown medially, yellow to pale brown laterally, liaceae), det. Michael Lovave (LAE herbarium, sometimes with reddish tinge, longitudinal stripe PNG), 3♂ (00271386–00271388), 1♀ (00271432) dark brown; anterior side yellow to brown; frons (ZISP). MALAYSIA: Sabah: West Coast Resi- brown, areas below antennal fossae whitish yel- dency, Ranau, 8 mi N Poring Hot Springs, low, clypeus yellow to pale brown, brown basally, 6.11407°N 116.72104°E, 500 m, 08 Oct 1958–11 sometimes with reddish tinge; tubercle around Oct 1958, T.C. Maa, 1♀ (00042310) (BPBM). antennal fossa whitish yellow with pale brown Sarawak: Bau, 1.4167°N 110.15°E, 29 Aug 1958– markings; lateral side whitish yellow to pale 30 Aug 1958, T.C. Maa, 1♂ (00042341), 1♀ brown, buccula, mandibular and maxillary plates (00042311) (BPBM). Merirai Valley nr. Kapit, whitish yellow; ventral side whitish yellow to 1.91009°N 113.63583°E, 01 Jul 1958–06 Jul 1958, pale brown, sometimes brown anteriorly. Eye T.C. Maa, 1♂ (00021523) (BPBM). Rajang pale brown. Labium: Segments I–II whitish yel- (Rejang) Delta, Sarikei Dist., 2.42873°N low, segment III whitish yellow, yellow ventrally, 111.44251°E, 15 Jul 1958–26 Jul 1958, T.C. Maa, segment IV yellow. Antenna: Segment I yellow to 1♂ (00021526), 1♀ (00021526) (BPBM); 15 Jul pale brown; segment II pale brown to brown, 1958–25 Jul 1958, T.C. Maa, 4♂ (00042337, sometimes with reddish tinge, segments III–IV 00021519–00021521), 2♀ (00021524, 00021525) brown to dark brown. Thorax: Anterior part of (BPBM). Sadong, Kampong Tapuh, 1.50361°N pronotum and punctures between anterior and 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 99 posterior parts brown, anterior margin dark slightly surpassing it; segments I and II strongly brown, posterior part of pronotum whitish yel- reduced; combined subequal to half of segment low with brown marking near posterior angle; III; segment I as long as wide; segment II slightly scutellum and mesoscutum whitish yellow to longer than wide, its dorsal surface elongate pos- pale brown; mesoscutum sometimes somewhat teriorly; segment III slightly longer than ventral darker than scutellum; punctures between scu- side of head; segment IV twice as long as seg- tellum and mesoscutum pale brown; thoracic ment III. Antenna: Segment I cylindrical (as in pleura brown to dark brown; scent gland evapo- Namyatova et al., 2016: fig. 8A), ca. 1.2–1.6× as rative area uniformly whitish yellow. Hemelytron: long as head width, ca. 0.9–1.1× as long as pro- Clavus translucent, its inner part pale brown to notum width; segment II ca. 1.9× as long as head brown with margins brown, sometimes with red- width, ca. 1.3× as long as pronotum width; seg- dish tinge, outer part of clavus colorless, pale ment III slightly longer than segment II; segment brown to brown apically; corium translucent IV ca. 0.25× as long as segment IV. Thorax: Ante- with marking along inner margin of corium rior and posterior parts of pronotum subequal in brown, narrow, not reaching R+M; embolium length; collar delimited; posterior part slightly opaque, whitish yellow, pale brown or reddish upraised; posterior margin straight or slightly apically, with brown margins, opaque; cuneus concave, pronotum ca. 1.2–1.3× as wide as long mostly opaque, reddish with colorless and trans- and ca. 1.4–1.5× as wide as head; mesoscutum lucent outer part and brown outer margin; mem- exposed. Hemelytron: Marking along inner mar- brane translucent with grayish tinge; membrane gin of corium almost flat; inner margin of cuneus cell pale brown veins. Legs: Coxae whitish yel- convex (as in Namyatova et al., 2016: fig. 13E), low; femora whitish yellow basally, yellow api- outer margin of cuneus almost ca. 2.5× as long cally, tibiae mostly yellow or pale brown, often as base. Abdomen: Genital capsule rotated left at whitish yellow apically, fore- and middle tarsi right angle relative to the rest of abdomen. Geni- whitish yellow to pale brown, hind tarsus whitish talia: Genital capsule (fig. 14R) ca. 1.5× as long yellow to yellow. Abdomen: Whitish yellow ven- as wide; ventral wall ca. 1.5× as long as dorsal trally and pale brown with reddish tinge dorsally. wall, its posterior margin smooth, semioval, SURFACE AND VESTITURE: Corium smooth, without outgrowth, not curved, its apex inclined with shallow and scarce punctures. Dorsum, leftward; sides of genital capsule not modified; antennal segment I and femora with suberect right paramere socket slightly angulate and left setae shorter than antennal segment II diameter; socket rounded; distance between paramere abdomen clothed with suberect setae of different sockets ca. 0.7× as long as genital capsule width length. STRUCTURE AND MEASUREMENTS: at base. Right paramere (fig. 12I) distinctly Body ca. 4.4–4.6× as long as pronotum width. curved in apical half; apex slightly concave; Head: Depression, delimiting occipital region medial part slightly wider than basal part, bear- distinct dorsally and laterally (as in Namyatova ing setae, with outer margin concave and inner et al., 2016: fig. 4E); distance between depression margin convex; outer angle distinct; inner angle and pronotum distinctly shorter than eye diam- rounded, without setae; basal part of paramere eter; longitudinal sulcus on dorsal surface longer ca. 0.15–2× as long as rest of paramere. Left than eye diameter; distance from eye to prono- paramere (fig. 12J) widened, only slightly curved; tum slightly longer than eye diameter, not swol- apical part straight, with toothlike outgrowth on len laterally (as in Namyatova et al., 2016: fig. posterior side medially (as in fig. 11G) and with- 4E); vertex ca. 1.2–1.3× as wide as eye, upraised out outgrowth on dorsal surface; middle part of (as in Namyatova et al., 2016: fig. 6D). Labium paramere widened, without swelling or out- (as in Namyatova et al., 2016: figs. 6D, 9C): growth; setae only on middle part near outer Reaching posterior margin of metasternum or margin. Aedeagus (general view as in Namyatova 100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 et al., 2016: fig. 22I) conjunctiva weakly sclero- ever, it has near identical vesical spicules, with F. tized; secondary gonopore placed at base of javanus (cf. fig. 8N with 8O), sharing spicules A, vesica in repose; sclerotization of ductus seminis B, and C, with all of them similar in shape. These around secondary gonopore shorter than wide; species also have similar body size and a red vesica with five spicules, including spicules A, B, cuneus, but F. javanus differs by the anterior part and C (fig. 8O). of the pronotum being yellow to pale brown, the Female. Total length 3.2–3.5. COLORATION inner part of the clavus is mostly whitish yellow (fig. 5): Head: Similar to male, but dorsal surface to yellow, with reddish or pale brown margins, sometimes yellow to pale brown with reddish and the labium reaches the posterior margin of tinge, brown anteriorly; frons yellow to brown, the mesosternum or slightly surpasses it. sometimes with reddish tinge; clypeus yellow Material examined: Holotype: AUSTRA- sometimes with reddish tinge, sometimes brown LIA: Territory of Christmas Island: Grants basally; ventral side whitish yellow to pale brown. Well. Christmas Is., 10.48333°S 105.65°E, 27 Apr Labium and antenna: As in male. Thorax: Similar 1989, J.C. Cardale, 1♂ (00033781) (ANIC). to male, but anterior part of pronotum some- Paratypes: AUSTRALIA: Territory of Christ- times pale brown with reddish tinge and brown mas Island: Grants Well. Christmas Is., anterior margin; thoracic pleura yellow to brown, 10.48333°S 105.65°E, 15 Apr 1989, J.C. Cardale, embolium translucent. Abdomen: Whitish yellow 2♂ (00033781, 00033783), 1♀ (00033784) with segment X and dorsal surface red. SUR- (ANIC); 27 Apr 1989, J.C. Cardale, 4♀ FACE AND VESTITURE: As in male. STRUC- (00033785–00033788) (ANIC). TURE AND MEASUREMENTS: Structure as in male; body ca. 4.1–4.4× as long as pronotum width; vertex ca. 1.8–1.9× as wide as eye diame- Felisacus linae, sp. nov. ter; antennal segment I ca. 1.4–1.6× as long as head width, ca. 0.9–1.0× as long as pronotum Figures 5, 8P, 12K, L, 14S, 18 width; segment II ca. 1.7–1.9× as long as head Diagnosis: Recognized by the following width, ca. 1.1–1.3× as long as pronotum width; combination of characters: dorsal surface of head pronotum ca. 1.2–1.3× as wide as long and ca. whitish yellow to pale brown; pronotum mostly 1.5–1.7× as wide as head. Genitalia (as in Namya yellow to pale brown with markings posteriorly tova et al., 2016: fig. 23F, G): Dorsal labiate plate and with collar darker than remainder of prono- wider than distance between apodemes of sec- tum; cuneus with yellow tinge or with yellow to ond valvula; mostly smooth, without distinct pale brown inner margin; cylindrical antennal striations, with semicircular sclerite and distinct segment I (as in Namyatova et al., 2016: fig. 8A), sclerotized rings laterally; lateral oviducts placed transverse depression on head extending later- almost medially, very close to each other, sper- ally; vertex upraised; labium reaching posterior mathecal gland placed between lateral oviducts; margin of mesosternum; antennal segment I ca. dorsal labiate plate with distinct tubercles, with- 1.7–1.9× as long as head width; cuneus ca. 3× as out membranous lobe medially. long as its base; apical part of right paramere Distribution: Christmas Is. (fig. 18). only slightly concave, medial part of right para- Host plants: Unknown. mere distinctly wider than basal part, shorter Etymology: The species is named after Chris- than basal and apical part combined (fig. 12K); tine Lambkin, curator of the Entomology in vesica with six spicules, including spicules A, B, Queensland Museum (Brisbane), who has assisted and C, spicule B shortened (fig. 8P). us in this work including the loan of material. Description: Male. Total length 3.7–3.9. Discussion: Felisacus lambkinae is not very COLORATION (fig. 5): Head: Mostly whitish similar in coloration to any other Felisacus. How- yellow to pale brown, often with brown or red 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 101 markings; clypeus sometimes brown. Eye dark with shallow, scarce punctures. Body clothed brown to black, often with reddish tinge. with pale simple setae; dorsum, antennal seg- Labium: Whitish yellow to yellow, segment III ment I clothed with setae subequal to antennal with pale brown or reddish stripe ventrally. segment II diameter; femora clothed with Antenna: Segment I yellow to paler brown, suberect setae shorter than antennal segment II sometimes with reddish tinge, somewhat paler diameter; abdomen clothed with short suberect basally; segment II pale brown to dark brown, setae. STRUCTURE AND MEASUREMENTS: often with reddish tinge; segments III–IV Body ca. 4.3–5.1× as long as pronotum width. brown to dark brown. Thorax: Pronotum mostly Head: Depression delimiting occipital region whitish yellow to pale brown, rarely with red- present dorsally and laterally (as in Namyatova dish longitudinal stripe laterally; collar often et al., 2016: fig. 4E); distance between depres- pale brown with anterior margin dark brown or sion and pronotum distinctly shorter than eye uniformly dark brown; punctures between ante- diameter; longitudinal sulcus on dorsal surface rior and posterior parts pale brown; posterior of head as long as eye diameter; distance from margin of pronotum with pair of brown to dark eye to pronotum distinctly longer than eye brown making near humeral angles, sometimes diameter, not swollen laterally (as in Namyatova posterior margin pale brown to brown between et al., 2016: fig. 4E); vertex ca. 1.3–1.8× as wide markings; scutellum and mesoscutum whitish as eye, upraised (as in Namyatova et al., 2016: yellow to pale brown; scutellum sometimes with fig. 6D). Labium (as in Namyatova et al., 2016: pale brown to brown marking or stripe medially figs. 6D, 9C): Reaching posterior margin of or apically, punctures between mesoscutum and mesosternum or slightly surpassing it; segments scutellum often pale brown; thoracic pleura I and II strongly reduced, combined shorter whitish yellow to pale brown; scent gland evap- than half of segment III; segment I shorter than orative area whitish yellow, paler than pleura, wide; segment II as long as wide, elongate dor- often reddish apically. Hemelytron: Mostly sally; segment III as long as ventral side of head; translucent, colorless; inner part of clavus segment IV ca. 1.5× as long as segment III. sometimes opaque, yellow to brown with brown Antenna: Segment I cylindrical (as in Namya- margins; outer part of clavus often with pale tova et al., 2016: fig. 8A), ca. 1.7–1.9× as long as brown to brown apex; area along inner margin head width, ca. 1.1–1.2× as long as pronotum of corium brown, often somewhat darker ante- width; segment II ca. 2.1–2.4× as long as head riorly; embolium yellow to pale brown apically width, ca. 1.4–1.5× as long as pronotum width; with brown margins; cuneus with colorless segment III slightly longer than segment II; seg- outer part and yellow to pale brown inner part ment IV ca. 0.3× as long as segment III. Thorax: with pale brown or brown margins, sometimes Anterior part of pronotum distinctly shorter with yellow tinge and pale brown margins; than posterior part, collar delimited; posterior membrane with yellow cell. Legs: Coxae whitish part slightly upraised; posterior margin of pro- yellow; femora whitish yellow basally and yel- notum straight or slightly concave; pronotum low to pale brown apically, sometimes with red ca. 1.2–1.3× as wide as long and ca. 1.5–1.7× as stripe at least apically; tibiae yellow to pale wide as head; mesoscutum exposed or not brown, often paler apically, sometimes with red exposed. Hemelytron: Area along inner margin stripe at least apically; tarsi yellow to brown, of corium not swollen; inner margin of cuneus sometimes segments I and II yellowish and seg- convex (as in Namyatova et al., 2016: fig. 13E), ment II brown. Abdomen: Ventral and lateral outer margin of cuneus ca. 3× as long as base. sides whitish yellow to yellow with genital seg- Abdomen: Genital capsule rotated left at right ment yellow or red, dorsal surface red. SUR- angle relative to rest of abdomen. Genitalia: FACE AND VESTITURE: Corium smooth, Genital capsule (fig. 14S) ca. 1.5× as long as 102 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 wide; ventral wall almost twice as long as dorsal plate wider than distance between apodemes of wall, with posterior margin of ventral wall second valvula; mostly smooth, without distinct semioval, smooth, without outgrowth(s), its striations, with semicircular sclerite and distinct apex inclined rightward; sides of genital capsule sclerotized rings laterally; lateral oviducts placed not modified; right paramere socket slightly almost medially, very close to each other, sper- acute, left one rounded; distance between para- mathecal gland placed between lateral oviducts; mere sockets subequal to half of genital capsule dorsal labiate plate with distinct tubercles, with- width at base. Right paramere (fig. 12K) dis- out membranous lobe medially. tinctly curved in apical half; apex concave pos- Distribution: Portuguese Timor (fig. 18). teriorly; medial part only slighter wider than Host plants: Unknown. basal part, bearing setae, with outer margin Etymology: The species is named after Meiy- straight and inner margin convex; outer angle ing Lin (Institute of Zoology, Chinese Academy distinct, only slightly widened; inner angle of Sciences), who helped us with a loan of Feli- indistinct, not bearing setae; basal part of para- sacus specimens. mere ca. 0.15–0.2× as long as rest of paramere. Discussion: Felisacus linae is externally simi- Left paramere (fig. 12L) L-shaped; apical part lar to F. filicicola, F. ochraceus, and F. solomonicus not flattened, with toothlike outgrowth on pos- (figs. 5, 6). These three species differ from F. linae terior side medially (as in fig. 11G) and without in the collar the same color as the rest of the pro- outgrowth on dorsal surface; middle part wid- notum. Felisacus filicicola also differs in having ened, without swelling or outgrowth; setae only seven vesical spicules (fig. 8I). Felisacus ochraceus on middle part near outer margin. Aedeagus also can be separated by the following characters (general view as in Namyatova et al., in press: in the male genitalia, with the right paramere fig. 22I) conjunctiva weakly sclerotized; second- narrow medially, only slightly wider than the ary gonopore placed at base of vesica; scleroti- basal part, almost as long as basal and apical zation of ductus seminis at secondary gonopore parts combined (fig. 13C), and it has five vesical shorter than wide; vesica with six spicules, spicules, with spicule B elongate (fig. 9F). Feli- including spicules A, B, and C. Spicule A long sacus solomonicus is also similar to F. linae in and wide, spicule B shortened, swollen medi- coloration, but differs in smaller size, with the ally, spicule C distinctly moved posteriorly, long body length of males 3.1–3.4 mm and females and convolute basally (fig. 8P). 3.3–3.4 mm. It also has the right paramere dis- Female. Total length 3.8–4.2. COLORATION tinctly concave apically (fig. 13R) and has five (fig. 5): Mostly as similar to male, but scutellum vesical spicules, lacking spicules B and C (fig. sometimes mostly pale brown with yellow mar- 9L). gins or uniformly pale brown. Abdomen: Ventral Material examined: Holotype: EAST side whitish yellow to yellow, lateral and dorsal TIMOR: Estação Zootecnica and foot of Mundo surfaces reddish. SURFACE AND VESTITURE: Perdido nr Ossu, Timor Portugues, 8.73423°S As in male. STRUCTURE AND MEASURE- 126.35914°E, 15 Mar 1961, G.F. Gross, 1♂ MENTS: Structure as in male; body ca. 4.0–4.4× (00017820) (SAMA). Paratypes: EAST TIMOR: as long as pronotum width; vertex ca. 1.6–1.9× Esçãao Zootecnica and foot of Mundo Perdido as wide as eye; antennal segment I ca. 1.4–1.8× nr Ossu, Timor Portugues, 8.73423°S as long as head width, ca. 0.9–1.0× as long as 126.35914°E, 14 Mar 1961, G.F. Gross, 6♂ pronotum width; segment II ca. 1.8–2.1× as long (00017813–00017818), 1 sex unknown as head width, ca. 1.1–1.4× as long as pronotum (00017823) (AM); 15 Mar 1961, G.F. Gross, 2♂ width; pronotum ca. 1.2–1.4× as wide as long (00017819, 00017821), 1 sex unknown and ca. 1.4–1.8× as wide as head. Genitalia (figs. 00017822), 1 juvenile (00017811), 7♀ 23F, G in Namyatova et al., 2016): Dorsal labiate (00017824–00017830) (SAMA). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 103
Felisacus lindbergae, sp. nov. translucent, colorless, with brown margins and apex sometimes with brown tinge; membrane Figures 5, 10F, 12M, N, 14T, 16E, 19 translucent, gradually changing color from Diagnosis: Recognized by the following brown anteriorly and pale brown posteriorly, combination of characters: antennal segment II sometimes membrane pale brown anteriorly and brown, cuneus colorless, sometimes with brown colorless posteriorly; cell brown. Legs: Coxae tinge apically; femora with marking in apical half whitish yellow, femora whitish yellow basally and brown to dark brown; marking on corium dark yellow apically with apex and marking in apical brown, C-shaped, its anterior part reaching or half brown to dark brown, sometimes marking almost reaching R+M and only slightly inclined; in apical half very pale; foretibia mostly brown, antennal segment I widened (as in Namyatova whitish yellow to yellow apically, sometimes and Cassis, in press: fig. 8B), ventral wall of geni- brown apically and yellow apically; middle and tal capsule with tooth posteriorly (fig. 14T); right hind tibiae brown basally and whitish yellow api- paramere sickle shaped with inner angle bearing cally; tarsi mostly whitish yellow, sometimes seg- straight outgrowth (fig. 12M); left paramere with ments II and III of fore- and middle tarsi pale tooth apically and outgrowth on dorsal surface, brown to brown. Abdomen: Whitish yellow to but without outgrowth on inner margin (fig. yellow with brown dorsum. SURFACE AND 12N); half of ductus seminis sclerotized, sickle VESTITURE: Corium smooth, with shallow and shaped apically; vesica without sclerite (fig. 10F). scarce punctation. Dorsum and forefemora with Description: Male. Total length 3.4–4.1. setae subequal to or longer than antennal seg- COLORATION (fig. 5): Head: Mostly yellow to ment II diameter; antennal segment I, middle pale brown, dorsal surface sometimes with and hind femora clothed with suberect setae brown marking near eye, longitudinal sulcus mostly shorter than antennal segment II diame- sometimes brown, darker than dorsal surface, ter; abdomen clothed with short erect setae. anterior side and clypeus basally sometimes yel- STRUCTURE AND MEASUREMENTS: Body low, paler than other sides, clypeus apically and ca. 4.1–4.5× as long as pronotum width. Head: buccula brown. Eye dark brown with reddish Depression, delimiting occipital region, present tinge. Labium: Segments I–II dark brown, seg- only dorsally; distance between depression and ment III–IV yellow to brown, paler than segment pronotum distinctly longer than eye diameter; II, segment IV sometimes darker than segment longitudinal sulcus on dorsal surface of head III. Antenna: Segments I–II brown, segments shorter than eye diameter; distance from eye to III–IV yellow to pale brown. Thorax: Anterior pronotum distinctly longer than eye diameter, part of pronotum mostly yellow to pale brown, not swollen laterally; vertex ca. 1.4–1.6× as wide often brown medially, anterior margin brown, as eye, flat (Namyatova et al., 2016: fig. 6E). posterior part of pronotum uniformly brown; Labium: Almost reaching posterior margin of scutellum and mesoscutum brown to dark prosternum, but not surpassing it; segments I brown; thoracic pleura brown to dark brown; and II longer than wide, each of them subequal scent gland evaporative area uniformly whitish to half of segment III; dorsal surface of segment yellow. Hemelytron: Clavus opaque brown to II not elongate posteriorly (as in Namyatova et dark brown; corium and embolium translucent, al., 2016: fig. 6E); segment III distinctly shorter colorless; corium with C-shaped dark brown than ventral side of head; segment IV ca. 1.5× as marking, its anterior part of this marking reach- long as segment III. Antenna: Segment I swollen ing or almost reaching R+M vein; posterior part in basal portion (as in Namyatova et al., 2016: of C-shaped marking reaching costal margin; fig. 8B), ca. 1.2–1.3× as long as head width, ca. embolium with brown outer margin and color- 0.8–0.9× as long as pronotum width; segment II less inner margin and dark brown apex; cuneus ca. 1.5–1.8× as long as head width, ca. 1.1–1.3× 104 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 as long as pronotum width; segment III slightly ture as in male; body ca. 4.1–4.3× as long as pro- longer than segment II and twice as long as seg- notum width; vertex ca. 1.5–1.8× as wide as eye ment IV. Thorax: Anterior part of pronotum diameter; antennal segment I ca. 1.1–1.2× as shorter than posterior part; collar delimited; pos- long as head width, ca. 0.7–0.8× as long as pro- terior part slightly upraised; posterior margin of notum width; segment II ca. 1.5–1.8× as long as pronotum straight or concave; pronotum ca. head width, ca. 1.0–1.1× as long as pronotum 1.0–1.3× as wide as long and ca. 1.3–1.5× as wide width; pronotum ca. 1.1–1.2× as wide as long as head; mesoscutum usually exposed, some- and ca. 1.5–1.7× as wide as head. Genitalia: Dor- times not exposed. Hemelytron: Area along inner sal labiate plate very small and transparent, as margin of corium swollen; inner margin of wide as distance with apodemes of second valvu- cuneus straight (as in Namyatova et al., 2016: fig. lae, without striations or membranous ridge 13F), outer margin of cuneus twice as long as its medially; semicircular sclerite and sclerotized width at base. Abdomen: Genital capsule rotated rings absent; lateral oviducts placed in near pos- left at small angle relative to rest of abdomen. terior margin very close to each; spermathecal Genitalia: Genital capsule (fig. 14T) ca. 1.5× as gland attached near anterior margin (fig. 16E); long as wide; ventral wall ca. 1.5× as long as dor- dorsal labiate plate without distinct tubercles, sal wall, with posterior margin of ventral wall without membranous lobe posteriorly (as in fig. truncate, not curved dorsally, with small tooth- 16B). like outgrowth from left side and with right side Distribution: Solomon Is. (fig. 19) folded; left side of genital capsule not widened; Host plants: Freycinetia sp. (Pandanaceae). paramere sockets distinctly angulate; distance Etymology: The species is named after between paramere sockets subequal to half of Gunvi Lindberg, the curatorial assistant in Swed- width of genital capsule base at base. Right para- ish Museum of Natural History (Stockholm) for mere (fig. 12M) sickle shaped; apex straight pos- the loan of type specimens. teriorly; medial part narrower than basal part, Discussion: Felisacus lindbergae is most simi- without setae, outer margin of medial part con- lar to F. longiceps, F. magnificus, and F. zuparkoi vex and inner margin concave, without swelling; in coloration and structure (figs. 5–7). These outer angle absent; inner angle present, with three species differ by the labium reaching the straight outgrowth, bearing setae; basal part of posterior margin of the mesosternum, the out- paramere ca. 0.7× as long as rest of paramere. growth on the inner angle of the right paramere Left paramere (fig. 12N) curved medially; apical is curved (figs. 12M, 12Q, W, 13AD), and the part not flattened, with toothlike outgrowth on medial part of the left paramere has an out- posterior side apically and with outgrowth on growth (cf. figs. 12N, 12R, X, 13AE).Felisacus dorsal surface; inner margin of paramere slightly magnificus can also be separated by the right swollen dorsally, bearing setae. Aedeagus (gen- paramere straight (fig. 12W) and the apical part eral view as in Namyatova et al., 2016: fig. 22M) of the left paramere broad (fig. 12X). Felisacus vesica without spicules; apical half of ductus zuparkoi differs from the above species by the seminis sclerotized, secondary gonopore placed femora without markings apically and the pres- near phallotheca mouth in repose; ductus semi- ence of a vesical spicule (fig. 10P). nis hooked apically (fig. 10F). Material examined: Holotype: SOLOMON Female. Total length 3.7–4.4. COLORATION ISLANDS: New Georgia Islands: Gizo, Gizo Is., (fig. 5): Similar to male, antennal segments III– 8.1°S 156.85°E, 100 m, Nov 1975, N.L.H. Krauss, IV yellow, paler than segment II, sometimes seg- 1♂ (00043172) (BPBM). Paratypes: SOLOMON ment IV pale brown, darker than segment III. ISLANDS: Malaita: Auki, 8.767°S 160.7°E, 100 m, SURFACE AND VESTITURE: As in male. Feb 1985, N.L.H. Krauss, 2♂ (00043164, 00043165) STRUCTURE AND MEASUREMENTS: Struc- (BPBM). New Georgia Islands: Gizo, Gizo Is., 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 105
8.1°S 156.85°E, 100 m, Nov 1975, N.L.H. Krauss, brown with collar pale brown and anterior mar- 8♂ (00043166–00043171, 00043173, 00043174), gin dark brown; posterior part of pronotum 2♀ (00043175, 00043176) (BPBM). Western Prov- whitish yellow to yellow, with small brown ince: New Georgia Islands: Munda, New Georgia marking at posterior margin medially and Is., 8.31666°S 157.25°E, 50 m, Nov 1980, N.L.H. brown marking near posterior angle; scutellum Krauss, 1♀ (00043178) (BPBM). New Georgia and mesoscutum whitish yellow, with brown Island, Munda, 8.31666°S 157.25°E, 20 Jul 1959, J.L. longitudinal stripe medially, rarely uniformly Gressitt, Freycinetia sp. (Pandaceae), 1♂ (00043163) whitish yellow; thoracic pleura pale brown to (BPBM). Gizo, Kolombangara Is., Gizo, 7.93799°S dark brown; scent gland evaporative area pale 157.09339°E, 70 m, Dec 1980, N.L.H. Krauss, 2♀ brown to brown with reddish apex. Hemelytron: (00043177–00043179) (BPBM). Mostly colorless, translucent; inner part of cla- vus with orange, reddish, or pale brown mar- Felisacus liui, sp. nov. gins; brown marking along inner margin of Figures 5, 8Q, 12O, P, 14U, 19 corium narrow, not reaching R+M; embolium with pale brown margins; cuneus with pale Diagnosis: Recognized by the following brown, orange, or reddish marking along inner combination of characters: small body, length in margin and pale brown margins, rarely cuneus male 2.6–2.8, in female 2.9; dark coloration with colorless with yellow margins. Legs: Coxae dorsal surface of head and anterior part of pro- whitish yellow; femora whitish yellow to yellow notum mostly brown, posterior part of prono- with reddish apices; tibiae reddish basally and tum pale brown with brown marking posteriorly, whitish yellow to yellowish apically; foretibia marking along inner margin of corium brown, sometimes uniformly reddish; tarsi whitish yel- not reaching R+M; and cuneus with narrow red low to yellow. Abdomen: Yellow with red or stripe along inner margin; cylindrical antennal brown lateral sides, apex and dorsal surface segment I (as in Namyatova et al., 2016: fig. 8A), sometimes uniformly yellow with reddish tinge. transverse depression on head extending later- SURFACE AND VESTITURE: Corium smooth, ally, vertex upraised (as in Namyatova et al., often with shallow scarce punctures. Dorsum 2016: fig. 6D); antennal segment I ca. 1.2–1.5× as with suberect setae mostly shorter than or sub- long as head width; labium reaching posterior equal to antennal segment II diameter; antennal margin of mesosternum or slightly surpassing it; segment I and femora with suberect setae middle part of right paramere almost as wide as shorter than antennal segment II diameter; basal part, shorter than basal and apical parts abdomen clothed with suberect mostly short combined (fig. 12O); vesica with six spicules, simple setae. STRUCTURE AND MEASURE- none of which can be homologized with the spic- MENTS: Body ca. 4.5–4.8× as long as pronotum ules of other species (fig. 8Q) width. Head: Transverse depression delimiting Description: Male. Total length 2.6–2.8. occipital region present dorsally and laterally COLORATION (fig. 5): Head: Varying from (as in Namyatova et al., 2016: fig. 4E); distance dark brown to pale brown; buccula, clypeus, between depression and pronotum distinctly and tubercle around antennal fossa yellow, shorter than eye diameter; longitudinal sulcus mandibular and maxillary plates yellow to pale on dorsal surface longer than eye diameter; dis- brown. Eye brown with reddish tinge. Labium: tance from eye to pronotum slightly longer than Yellow to pale brown, segment III often brown eye diameter, not swollen laterally (as in Namy- or reddish ventrally and basally. Antenna: Seg- atova et al., 2016: fig. 4E); vertex ca. 2.0–2.3× as ment I reddish with yellow base, segment II wide as eye, upraised (as in Namyatova et al., reddish brown or brown, segments III–IV 2016: fig. 6D). Labium (as in Namyatova et al., brown. Thorax: Anterior part or pronotum 2016: figs. 6D, 9C): Reaching posterior margin 106 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 of mesosternum or slightly surpassing it; seg- et al., 2016: fig. 22I) conjunctiva weakly sclero- ments I and II strongly shortened, combined tized; secondary gonopore placed at base of shorter than half of segment III; segment II vesica in repose; sclerotization of ductus semi- slightly longer than wide, its dorsal surface nis around secondary gonopore shorter than elongate posteriorly; segment III slightly longer wide; vesica with six spicules, none of which than ventral side of head; segment IV twice as can be homologized with the spicules of other long as segment III. Antenna: Segment I cylin- species (fig. 8Q). drical (as in Namyatova et al., 2016: fig. 8A), ca. Female. Total length 2.9. COLORATION 1.2–1.4× as long as head width, ca. 0.9–1.1× as (fig. 5): Similar to male, but scutellum and long as pronotum width; segment II ca. 1.6– mesoscutum whitish yellow with brown longi- 1.8× as long as head width, ca. 1.2–1.4× as long tudinal stripe, scent gland evaporative area as pronotum width; segment III subequal to brownish with reddish apex; clavus sometimes segment II. Thorax: Anterior part of pronotum yellowish and opaque, cuneus whitish and only slightly shorter than posterior part; collar translucent with reddish inner part and pale delimited; posterior part slightly upraised; pos- brown margins. SURFACE AND VESTITURE: terior angles of pronotum not delimited with As in male. STRUCTURE AND MEASURE- depression; posterior margin concave, prono- MENTS: Structure as in male; body ca. 4.5–4.8× tum ca. 1.1–1.2× as wide as long and ca. 1.2– as long as pronotum width; vertex ca. 1.7–2.0× 1.4× as wide as head; mesoscutum exposed. as wide as eye; antennal segment I ca. 1.4–1.5× Hemelytron: Area along inner margin of corium as long as head width, ca. 0.9–1.1× as long as almost flat; inner margin of cuneus convex (as pronotum width; segment II ca. 1.8× as long as in Namyatova et al., 2016: fig. 13E), outer mar- head width, ca. 1.2–1.3× as long as pronotum gin of cuneus almost ca. 3× as long as base. width; pronotum ca. 1.1–1.2× as wide as long Abdomen: Genital capsule rotated left at right and ca. 1.4–1.5× as wide as head. Genitalia (as angle relative to the rest of abdomen. Genitalia: in Namyatova et al., 2016: fig. 23F, G): Dorsal Genital capsule (fig. 14U) ca. 1.5× as long as labiate plate wider than distance between wide; ventral wall ca. 1.5× as long as dorsal apodemes of second valvula; mostly smooth, wall, its posterior margin smooth, semioval, without distinct striations, with semicircular without outgrowth, not curved dorsally; its sclerite and distinct sclerotized rings laterally; apex inclined leftward; sides of genital capsule lateral oviducts placed almost medially, very not modified; margins of paramere sockets close to each other, spermathecal gland placed rounded; distance between paramere sockets between lateral oviducts; dorsal labiate plate subequal to half of genital capsule base. Right with distinct tubercles, without membranous paramere (fig. 12O) distinctly curved in apical lobe medially. half; apex straight; medial part slightly wider Distribution: Solomon Islands (Kolomban- than basal part, bearing setae, with outer mar- gara Is., Gizo Is., New Georgia Is.) (fig. 19). gin concave and inner margin convex; outer Host plants: Unknown. angle distinct, slightly swollen; inner angle Etymology: The species is named after Guo- indistinct, without setae; basal part of paramere Qing Liu, who loaned us the material from the ca. 0.2–0.25× as long as rest of paramere. Left collection of Nankai University (China). paramere (fig. 12P) L-shaped; apical part not Discussion: Feliscaus liui is most similar flattened, with tooth on posterior side medially externally and in dark coloration to F. lambkinae (as in fig. 11G) and without outgrowth on dor- and F. minutus. However, F. lambkinae and F. sal surface; middle part widened, without swell- minutus have a larger red marking on the cuneus ing or outgrowth; setae only on middle part (figs. 5, 6). Felisacus minutus is also recognized near outer margin. Aedeagus (as in Namyatova by the posterior part of the pronotum being 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 107 brown laterally. Felisacus lambkinae and F. minu- tubercle around antennal fossa yellow, often buc- tus also differ from F. liui in the shape of vesical cula and apex of clypeus and rarely lateral side spicules (cf. figs. 8Q, O, 9D). behind eye brown. Eye brown with reddish tinge. Material examined: Holotype: SOLOMON Labium: Segment I yellow to pale brown; segment ISLANDS: Western Province: Gizo, Kolomban- II pale brown; segment III yellow to pale brown; gara Is., Gizo, 7.93799°S 157.09339°E, 70 m, Dec segment IV yellow, rarely labium uniformly yel- 1980, N.L.H. Krauss, 1♂ (00043238) (BPBM). low to pale brown. Antenna: Segment I brown or Paratypes: SOLOMON ISLANDS: Western reddish brown; segment II yellow to dark brown, Province: New Georgia Islands: Munda, New segments III–IV pale brown. Thorax: Anterior Georgia Is., 8.31666°S 157.25°E, 50 m, Nov 1980, part of pronotum yellow to pale brown, some- N.L.H. Krauss, 1♀ (00043240) (BPBM). Gizo, times with brown anterior margin; posterior part Kolombangara Is., Gizo, 7.93799°S 157.09339°E, of pronotum brown, rarely yellow or pale brown; 70 m, Dec 1980, N.L.H. Krauss, 4♂ (00043234– scutellum and mesoscutum pale brown to brown; 00043237), 1♀ (00043239) (BPBM). thoracic pleura pale brown to brown, rarely yel- low, scent gland evaporative area whitish yellow. Hemelytron: Clavus opaque, uniformly brown to Felisacus longiceps Poppius, 1915 dark brown, rarely pale brown; corium, embo- Figures 5, 10H, 12Q, R, 14V, 19 lium, cuneus, and membrane mostly translucent, Felisacus longiceps Poppius, 1915b: 55 (original colorless; corium with pale brown anterior angles description). and C-shaped dark brown marking, reaching Felisacus okinawanus Miyamoto, 1965: 166; new R+M anteriorly and posteriorly, its anterior part synonymy. not inclined posteriorly; embolium pale brown basally and brown apically, with pale brown or Diagnosis: Recognized among congeners by yellow margins; cuneus yellow, red, or brown api- the following combination of characters: pres- cally, with reddish brown or yellow outer margin ence of dark brown C-shaped marking on and colorless inner margin; membrane gradually corium, with anterior part reaching R+M and changing color from pale brown to brownish not inclined posteriorly (fig. 5); antennal seg- basally and grayish apically; membrane cell pale ment I widened (as in Namyatova and Cassis, in brown to brown. Legs: Coxae whitish yellow; fem- press: fig. 8B), red, reddish brown, or brown; ora whitish yellow basally and brown apically, antennal segment II often mostly yellow; vertex often with reddish tinge apically, sometimes with flat and not widened laterally; labium reaching brown marking at apical part, rarely fore- and posterior margin of mesosternum; dark band in middle femora whitish yellow to yellow, with red- apical part of fore- and middle femora usually dish markings apically, sometimes with brown absent; right paramere sickle shaped with out- band on apical half of hind femur; basal part of growth on inner angle slightly curved (fig. 12Q); tibiae yellow to brown, often with reddish tinge, apical part of left paramere narrow, with tooth- apical part whitish yellow, rarely tibiae uniformly like outgrowth apically and outgrowth on dorsal whitish yellow; tarsi uniformly whitish yellow. surface; medial part of left paramere with nar- Abdomen: Whitish yellow to pale brown ventrally row outgrowth (fig. 12R); apical half of ductus and laterally, reddish apically and brown dorsally. seminis sclerotized, hooked; vesica without SURFACE AND VESTITURE: Corium smooth, sclerite (fig. 10H). with shallow and scarce punctation. Dorsum and Redescription: Male. Total length 3.0–3.4. femora clothed with setae subequal to or longer COLORATION (fig. 5): Head: Mainly pale brown, than antennal segment II diameter; antennal seg- sometimes with reddish markings; anterior side, ment I clothed with suberect setae shorter than clypeus, mandibular and maxillary plates, and antennal segment II diameter; abdomen clothed 108 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 with short erect setae. STRUCTURE AND MEA- rower than basal part, without setae, outer margin SUREMENTS: Body ca. 3.9–4.2× as long as pro- of medial part convex, inner margin concave, notum width. Head: Transverse depression without swelling; outer angle absent; inner angle delimiting occipital region present only dorsally; present with curved outgrowth bearing setae; distance between depression and pronotum basal part of paramere slightly shorter than rest of slightly longer than eye diameter; longitudinal sul- paramere. Left paramere (fig. 12R) apical part not cus on dorsal surface of head shorter than eye flattened, with toothlike outgrowth apically and diameter; distance from eye to pronotum longer with outgrowth on dorsal surface; inner margin than eye diameter, not swollen; vertex ca. 1.9–2.5× with narrow outgrowth, with setae near out- as wide as eye, flat (Namyatova et al., 2016: fig. growth. Aedeagus (general view as in Namyatova 6E). Labium: Reaching posterior margin of meso- et al., 2016: fig. 22M) vesica without spicules; api- sternum or slightly surpassing it; segments I and cal half of ductus seminis sclerotized, hooked api- II shorter than wide, combined subequal to half of cally, secondary gonopore placed near phallotheca segment III; dorsal surface of segment II not elon- mouth in repose (fig. 10H). gate (as in Namyatova et al., 2016: fig. 6E); seg- Female. Total length 3.5. COLORATION (fig. ment III shorter than ventral side of head; segment 5): Head: Similar to male, often with reddish IV ca. 1.5× as long as segment III. Antenna: Seg- markings on dorsal surface, frons, mandibular ment I slightly longer than head width, swollen and maxillary plates, and clypeus; antennal seg- basally (as in Namyatova et al., 2016: fig. 8B), ca. ment II often yellow, rarely pale brown apically or 1.0–1.3× as long as head width, ca. 0.6–0.7× as uniformly pale brown. Thorax: Similar to male, long as pronotum width; segment II ca. 1.6–1.7× but posterior part of pronotum sometimes pale as long as head width, ca. 0.8–1.0× as long as pro- brown with brown posterior margin. Abdomen: notum width; segment III slightly longer than seg- As in male. SURFACE AND VESTITURE: As in ment II and twice as long as segment IV. Thorax: male. STRUCTURE AND MEASUREMENTS: Anterior part of pronotum shorter than posterior Structure as in male; body ca. 3.8–4.0× as long as part; collar delimited, posterior part slightly pronotum width; vertex ca. 1.8–2.2× as wide as upraised; posterior angles not delimited with eye; antennal segment I ca. 1.1–1.2× as long as depression; posterior margin of pronotum con- head width, ca. 0.6× as long as pronotum width; cave; pronotum ca. 1.3× as wide as long and ca. segment II ca. 1.2–1.6× as long as head width, ca. 1.7–2.0× as wide as head; mesoscutum usually 0.7–0.8× as long as pronotum width; pronotum exposed, sometimes not exposed. Hemelytron: ca. 1.2–1.4× as wide as long and ca. 1.8–2.0× as Area along inner margin of corium swollen; inner wide as head. Genitalia (as in figs. 16A, B): Dorsal margin of cuneus straight (as in Namyatova et al., labiate plate very small and transparent, as wide 2016: fig. 13F), outer margin of cuneus twice as as distance with apodemes of second valvulae, long as base. Abdomen: Angle between genital without striations or membranous ridge medi- capsule and rest of abdomen varying from very ally; semicircular sclerite and sclerotized rings small to almost right. Genitalia: Genital capsule absent; lateral oviducts placed in anterior part; (fig. 14V) twice as long as wide; ventral wall ca. spermathecal gland attached near anterior mar- 1.5× as long as dorsal wall, with posterior margin gin; dorsal labiate plate without distinct tubercles, of ventral wall not curved, with small toothlike without membranous lobe posteriorly. outgrowth; right side of genital capsule folded, left Distribution: Borneo, Japan (Ryukyu Is.), side of genital capsule not widened; paramere Vietnam (fig. 19). sockets distinctly acute; distance between para Host plants: Many specimens were col- mere sockets ca. 0.3× as long as genital capsule lected from Nephrolepis biserrata (Davalliaceae) width at base. Right paramere (fig. 12Q) sickle and Plesioneuron sp. (Thelypteridaceae) on shaped; apex slightly concave; medial part nar- Ambon and Sulawesi islands. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 109
Discussion: Felisacus capitatus, F. longi- 11R). Felisacus lindbergae can be separated from ceps, F. magnificus, F. okinawanus, and F. pul- F. longiceps by the labium almost reaching the chellus are very similar externally (see posterior margin of the prosternum, the brown discussion for F. magnificus for details). The antennal segment II, the presence of dark bands identification of the material belonging to this on the fore- and middle femora and the straight group of species was not straightforward, as outgrowth on the inner angle of the right para- the types of F. longiceps, F. magnificus, and F. mere (fig. 12M). pulchellus are either females or had lost geni- Material examined: Holotype (Felisacus talia, and we could not examine the types of F. longiceps). TAIWAN: Chip-Chip, no date pro- capitatus and F. okinawanus from Kyushu Uni- vided, unknown collector, 1♀ (00018972) versity, where they are preserved according to (HNHM). Additional material: CHINA: Guan- the initial description (Miyamoto, 1965). dong: Mt Dinghu [Dinghu Shan, Dinhushan], We divided all the material examined for the 23.1751°N 112.5533°E, 28 Nov 1959, Rodendorf, above five taxa into three species, based on the 1♂ (00271521) (ZISP). INDONESIA: Maluku: structure of the male genitalia, particularly for Ambon Island nr Ambon City, 3.60706°S that of the parameres. We identified one of them 128.28328°E, 23 Aug 2012, F. Konstantinov, as Felisacus magnificus (see its discussion for Nephrolepis biserrata (Sw.) Schott (Davalliaceae), details). The second species is paler, often having det. Michael Lovave (LAE herbarium, PNG), 5♂ a pale brown pronotum; in this species if the (00271434–00271438), 2♀ (00271439, 00271440) posterior part of the pronotum is dark brown, (ZISP). West Nusa Tenggara: Lombok Island, then antennal segment I and the legs are almost Senaru, Panorama track to Sendang Gila Water- uniformly yellow. This pattern of coloration is fall, 8.30111°S 116.40833°E, 28 Aug 2012, F. Kon- similar to F. longiceps. The parameres and color- stantinov, 13♂ (00271441–00271453), 1 sex ation of this species are also similar to F. oki- unknown (00271351), 8♀ (00271454–00271461) nawanus (figs 12Q, R; Miyamoto, 1965: figs. (ZISP); 30 Aug 2012, F. Konstantinov, Nephrole- 43–46). On the basis of these observations, we pis biserrata (Sw.) Schott (Davalliaceae), det. synonymized F. okinawanus with F. longiceps. Michael Lovave (LAE herbarium, PNG), 9♂ The third species was unlike any other among (00271462–00271470), 2 sex unknown the described taxa, and we describe it as a new (00271471, 00271472), 9♀ (00271473–00271481) species, F. zuparkoi. (ZISP). Lombok Island, Senaru, track to Tiu Felisacus magnificus and F. zuparkoi differ Kelep Waterfall, 8.30111°S 116.40833°E, 31 Aug from F. longiceps externally by the antennal seg- 2012, F. Konstantinov, Nephrolepis biserrata (Sw.) ment II, and the posterior part of pronotum dark Schott (Davalliaceae), 4♂ (00271482–00271484, brown, whereas both species have dark bands on 00271424) Plesioneuron sp. (Thelypteridaceae), the fore- and middle femora. Felisacus magnifi- 1♂ (00386467) (ZISP). Lombok Island, nr cus also differs from F. longiceps by the straight Senaru, 8.31689°S 116.40381°E, 29 Aug 2012, F. right paramere (fig. 12W) and the broad left Konstantinov, 11♂ (00271485–00271495), 8♀ paramere (fig. 12X). F. zuparkoi differs in the (00271496–00271499, 00271501–00271504), 1 broad outgrowth on the inner margin of the left sex unknown (00271350) (ZISP). Lombok Island, paramere (fig. 13AE) and the presence of a vesi- nr Senaru, 8.31956°S 116.405°E, 31 Aug 2012, F. cal spicule (fig. 10P). Konstantinov, Nephrolepis biserrata (Sw.) Schott Felisacus longiceps is also similar to F. ceyloni- (Davalliaceae), 1♀ 00271500) Plesioneuron sp. cus and F. lindbergae. Felisacus ceylonicus differs (Thelypteridaceae), det. Michael Lovave (LAE from F. longiceps by its vertex broad laterally herbarium, PNG), 5♂ (00386617–00386621), behind the eye and the outgrowth on the medial 3♀ (00271515–00271517) (ZISP). JAPAN: Oki- part of the left paramere distinctly curved (fig. nawa: Ishigaki Is., Ryukyu Islands, 24.4°N 110 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
124.2°E, 01 Nov 1952, G.E. Bohart, 2♂ vesica with six spicules, including spicules B (00043262, 00043263) (BPBM); 10 Nov 1952–15 and E, spicule B short and widened (fig. 8R). Nov 1952, G.E. Bohart, 1♂ (00043261) (BPBM); Description: Male. Total length 3.5–4.0. 14 Oct 1999, Belokobylskij, 1♂ (00271505) COLORATION (fig. 5): Head: Mostly whitish yel- (ZISP). MALAYSIA: Sabah: Kalabakan, low to yellow, often with reddish tinge or red 4.4167°N 117.4833°E, 08 Nov 1958–15 Nov 1958, markings; clypeus often pale brown or red api- T.C. Maa, 1♂ 00043186) (BPBM). West Coast cally. Eye brown with reddish tinge or red. Labium: Residency [Coast Residency], Ranau, 6.08333°N Whitish yellow to yellow, segment II often reddish 116.5°E, 500 m, 08 Oct 1958–11 Oct 1958, T.C. ventrally. Antenna: Segment I yellow with reddish Maa, 2♂ (00043344, 00043345) (BPBM). PHIL- tinge; segment II yellow with pale brown or red- IPPINES: Laguna: Los Banos, 14.17086°N dish apex; segment III yellow basally and pale 121.24334°E, no date provided, Baker [no date], brown to brown apically; segment IV pale brown 1♀ (00018446) (MZH). TAIWAN: N Taiwan, to brown. Thorax: Pronotum whitish yellow to Nei-Hu, Taipei, 25.09074°N 121.55972°E, 28 Aug yellow, often pale brown or brown posteriorly, 1963, Hsien, 1♀ (00005965) (BPBM). VIET- punctures between anterior and posterior parts NAM: Ha Noi: 70 km NWW Ha Noi BaVi, yellow to pale brown; scutellum and mesoscutum 21.087°N 105.302°E, 22 Nov 1990, Belokobylskij, whitish yellow to yellow, often with reddish stripe 1♀ (00018444) (ZISP). Hòa Bình: Hoa Binh: between them; thoracic pleura yellow; scent gland Near Mai Chau [Mai Chon, Mai Chou], 20.667°N evaporative area whitish yellow, rarely green api- 105.084°E, 31 Oct 1990, Belokobylskij, 3♀ cally. Hemelytron: Mostly translucent, colorless; (00018436, 00018442), 5♂ (00018437, 00018439, inner part of clavus whitish yellow to yellow, 00018440, 00271520) (ZISP); 01 Nov 1990, Belo- sometimes opaque, often brown basally, with pale kobylskij, 1♂ (00018438) (ZISP); 02 Nov 1990, brown, brown, or red margins; area along inner Belokobylskij, 1♀ (00018443) (ZISP); 03 Nov margin of corium brown; embolium with yellow 1990, Belokobylskij, 2♂ (00386603) (ZISP); 04 to pale brown margins; cuneus with pale brown Nov 1990, Belokobylskij, 2♂ (00018441) (ZISP). outer margin; membrane with grayish tinge; membrane cell red. Legs: Coxae whitish yellow; Felisacus lordhowensis, sp. nov. femora whitish yellow, often yellow apically, some- times with reddish tinge; tibiae yellow, often whit- Figures 5, 8R, 12S, T, 14W, 19 ish yellow apically, sometimes with reddish tinge Diagnosis: Recognized by the following basally, rarely uniformly whitish yellow or yellow; combination of characters: cylindrical antennal tarsi pale brown. Abdomen: Whitish yellow, often segment I (as in Namyatova et al., 2016: fig. yellow or red dorsally, rarely greenish basally. 8A), vertex upraised (as in Namyatova et al., SURFACE AND VESTITURE: Corium smooth, 2016: fig. 6D); head, clavus, and pronotum with shallow scarce punctures. Dorsum; antennal whitish yellow to yellow, head without dark segment I and femora with suberect setae shorter marking dorsally, pronotum sometimes with than antennal segment II diameter; abdomen narrow pale brown to brown marking along clothed with suberect mostly short simple setae. posterior margin, clavus uniformly colored; STRUCTURE AND MEASUREMENTS: Body ca. cuneus mostly colorless; labium reaching or 4.6–4.7× as long as pronotum width. Head: surpassing posterior margin of metasternum; Depression delimiting occipital region present antennal segment II ca. 1.6–1.9× as long as head dorsally and laterally (as in Namyatova et al., width; cuneus 3× as long as wide; medial part 2016: fig. 4E); distance between depression and of right paramere shorter than apical and basal pronotum distinctly shorter than eye diameter; parts combined (fig. 12S); apical part of left longitudinal sulcus on dorsal surface longer than paramere ca. 3× as long as wide (fig. 12T), eye diameter; distance from eye to pronotum eye 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 111 longer than eye diameter, not swollen laterally (as 11G) and without outgrowth on dorsal surface; in Namyatova et al., 2016: fig. 4E); vertex ca. 2.0– middle part widened, without swelling or 2.4× as wide as eye, upraised (Namyatova et al., outgrowth(s); setae only on middle part near 2016: fig. 6D). Labium: Reaching or slightly sur- outer margin. Aedeagus (general view as in passing posterior margin of metasternum; seg- Namyatova et al., 2016: fig. 22I) conjunctiva ments I and II strongly reduced, combined shorter weakly sclerotized, secondary gonopore placed at than half of segment III; segment I shorter than base of vesica in repose; sclerotization of ductus wide (as in Namyatova et al., 2016: figs. 6D, 9C); seminis around secondary gonopore shorter than dorsal surface of segment II elongate posteriorly, wide; vesica with six spicules, including spicules B elongate dorsally; segment III slightly longer than and E (fig. 8R), spicule B short and widened. ventral side of head; segment IV ca. 1.5× as long Female. Total length 3.9–4.5. COLORATION as segment III. Antenna: Segment I cylindrical (as (fig. 5): Similar to male, scent gland evaporative in Namyatova et al., 2016: fig. 8A), ca. 1.6–1.9× as area always whitish yellow; abdomen uniformly long as head width, ca. 1.0–1.1× as long as prono- yellow, often red dorsally, sometimes only apical tum width; segment II ca. 2.0–2.3× as long as head part of dorsal surface red. SURFACE AND VES- width, ca. 1.3–1.4× as long as pronotum width; TITURE: As in male. STRUCTURE AND MEA- segment III subequal to or slightly longer than SUREMENTS: Structure as in male; body ca. segment II; segment IV ca. 0.25–0.3× as long as 4.2–4.8× as long as pronotum width; vertex ca. segment IV. Thorax: Anterior part of pronotum 1.8–2.1× as wide as eye diameter; antennal seg- slightly shorter than posterior part; collar delim- ment I ca. 1.6–2.1× as long as head width, ca. ited; posterior part slightly upraised; posterior 1.0–1.1× as long as pronotum width; segment II ca. margin concave; pronotum ca. 1.1–1.2× as wide as 2.0–2.3× as long as head width, ca. 1.2–1.5× as long and ca. 1.5–1.8× as wide as head; mesoscu- long as pronotum width; pronotum ca. 1.2× as tum exposed. Hemelytron: Area along inner mar- wide as long and ca. 1.5–1.8× as wide as head. gin of corium almost flat; inner margin of cuneus Genitalia (as in Namyatova et al., 2016: fig. 23F, G): convex (as in Namyatova et al., 2016: fig. 13E), Dorsal labiate plate wider than distance between outer margin of cuneus ca. 3× as long as base. apodemes of second valvula; mostly smooth, with- Abdomen: Genital capsule rotated left at right out distinct striations, with semicircular sclerite angle relative to rest of abdomen. Genitalia: Geni- and distinct sclerotized rings laterally; lateral ovi- tal capsule (fig. 14V) twice as long as wide; ventral ducts placed almost medially, very close to each wall ca. 1.5× as long as dorsal wall, its posterior other, spermathecal gland placed between lateral margin smooth, semioval, without outgrowth, oviducts; dorsal labiate plate with distinct tuber- rounded, not curved; sides of genital capsule not cles, without membranous lobe medially. modified; right paramere socket slightly acute, left Distribution: Lord Howe Is. (fig. 19). paramere socket rounded; distance between para- Host plants: Unknown. mere sockets subequal to half of genital capsule Etymology: The species is named after Lord width at base. Right paramere (fig. 12S) distinctly Howe Island, where it was collected. curved in apical half; apex slightly concave poste- Discussion: Felisacus lordhowensis is similar riorly; medial part only slighter wider than basal to F. caledonicus, F. elegantulus, F. schuhi, and F. part, bearing setae, with outer margin slightly tanna in pale coloration (figs. 5, 6). In contrast concave and inner margin convex; outer angle dis- to F. lordhowensis, F. caledonicus has the labium tinct, widened; inner angle rounded, without reaching the middle of the mesosternum, the setae; basal part of paramere ca. 0.15–0.2× as long collar often pale brown to brown laterally (fig. as rest of paramere. Left paramere (fig. 12T) 4), the apical part of the left paramere is broad, L-shaped; apical part not flattened, with toothlike twice as long as wide (fig. 11Q) and four vesical outgrowth on posterior side medially (as in fig. spicules, including spicules A and B are present 112 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
(fig. 8F). Felisacus elegantulus differs from F. (fig. 12U); inner margin of medial part of left lordhowensis by the labium reaching the poste- paramere with triangular-shaped swelling (fig. rior margin of the metasternum, the humeral 12V); apical half of ductus seminis sclerotized, angles of the pronotum are pale brown to dark straight apically (fig. 10I). brown, and the presence of five vesical spicules, Description: Male. Total length 3.8. COL- including spicule B (fig. 8H). Felisacus schuhi ORATION (fig. 5): Head: Dorsal and lateral sides can be separated from F. lordhowensis by the mostly pale brown, anterior and ventral surfaces inner part of the clavus mostly brown, the mostly yellow; dorsal surface posteriorly and labium slightly surpasses the middle of the marking on frons reddish, buccula pale brown mesosternum, and the vesica has five spicules, anteriorly, mandibular and maxillary plates yel- including spicules A, B, and C (fig. 9J). Felisacus low; clypeus yellow, with reddish tinge basally tanna is most similar to F. lordhowensis, and and brown apically; eye reddish brown. Labium: differs from it by having five vesical spicules, Segments I, III, and IV yellow, segment II pale including spicules A, B, and E. (fig. 9M) brown. Antenna: Segments I–II reddish brown, Material examined: Holotype: AUSTRA- segments III–IV brown. Thorax: Pronotum pale LIA: New South Wales: On SW spur of Mt brown, brown posteriorly; mesoscutum and scu- Lidgebird, Lord Howe Island, 31.56666°S tellum pale brown; thoracic pleura mostly pale 159.08333°E, 122m, 12 Sep 1975, Lambkin, 1♂ brown, mesopleura with brown marking; scent (00017833) (AM). Paratypes: AUSTRALIA: New gland evaporative area whitish yellow. Hemely- South Wales: Lord Howe Is., Old Settlement, tron: Mostly translucent; clavus opaque, brown; 31.519°S 159.05083°E, 21 Feb 1971, D.K. McAl- corium colorless, marking along inner margin of pine, 2♂ (00017836, 00017837) (AM). Lord corium brown, reaching R+M posteriorly and Howe Island, 31.552°S 159.081°E, A.M. Lea, 3♂ not extending toward R+M anteriorly; embolium (00038900), 6♀ (00038900) (SAMA). On SW colorless with brown margins; cuneus with yel- spur of Mt Lidgebird, Lord Howe Island, low tinge, with outer margin brown and inner 31.56666°S 159.08333°E, 122m, 12 Sep 1975, margin whitish yellow; membrane with brown- Lambkin, 3♂ (00017832, 00017834, 00017835), ish tinge; membrane cell brown. Legs: Coxae 6♀ (00017838–00017843) (AM). whitish yellow, femora whitish basally and yellow apically; tibiae and tarsi yellow, foretibia and tar- Felisacus luzonus, sp. nov. sus somewhat darker than middle and hind tib- iae and tarsi. SURFACE AND VESTITURE: Figures 5, 10I, 12U, V, 14X, 16F, 19 Corium smooth, with shallow and scarce punc- Diagnosis: Recognized by the following tation. Dorsum with setae subequal to or longer combination of characters: marking along inner than antennal segment II diameter; antennal seg- margin of corium brown, narrow, reaching R+M ment I clothed with suberect setae shorter than posteriorly and not extending toward R+M ante- antennal segment II diameter; forefemur clothed riorly; antennal segment I swollen medially (as with suberect setae longer than antennal seg- in Namyatova et al., 2016: fig. 8B), red; trans- ment II diameter. STRUCTURE AND MEA- verse depression delimiting occipital region pres- SUREMENTS: Body ca. 4.0× as long as pronotum ent only dorsally; dorsal surface of labial segment width. Head: Depression delimiting occipital I not elongate posteriorly (as in Namyatova et al., region present only dorsally (Namyatova et al., 2016: fig. 6E); labium reaching middle of meso- 2016: fig. 6E); distance between depression and sternum; ventral wall of genital capsule curved pronotum shorter than eye diameter; longitudi- apically; genital capsule not swollen from left- nal sulcus on dorsal surface of head shorter than hand side (fig. 14X); right paramere with outer eye diameter; distance from eye to pronotum and inner angles distinct, without outgrowths distinctly longer than eye diameter, not swollen 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 113 laterally; vertex ca. 1.5× as wide as eye, flat (as in without spicules; half of ductus seminis sclero- Namyatova et al. 2016: fig. 6E). Labium: Reach- tized apically, secondary gonopore placed near ing middle of mesosternum; segments I and II theca mouth in repose; ductus seminis straight slightly longer than wide, combined longer than apically (fig. 10I). half of segment III; dorsal surface of segment II Female. Total length 4.1. COLORATION: not elongate posteriorly (as in Namyatova et al., Head: Mostly as in male, abdomen whitish with 2016: fig. 6E); segment III shorter than ventral brown dorsum. SURFACE AND VESTITURE: side of head, segment IV ca. 1.5× as long as seg- As in male. STRUCTURE AND MEASURE- ment III. Antenna: Segment I slightly longer than MENTS: Structure as in male; body ca. 4.1× as head width, widened medially (as in Namyatova long as pronotum width; vertex ca. 1.4× as wide et al., 2016: fig. 8B), ca. 1.4× as long as head as eye; pronotum ca. 1.2× as wide as long and width, ca. 0.8× as long as pronotum width; seg- 1.9× as wide as head. Genitalia: Dorsal labiate ment II ca. 1.7× as long as head width, ca. 0.9× plate very small and transparent, as wide as dis- as long as pronotum width; segment III slightly tance between apodemes of second valvulae, longer than segment II; segment IV subequal to without striations or membranous ridge medi- half of segment III. Thorax: Anterior part shorter ally; semicircular sclerite and sclerotized rings than posterior part; collar delimited; posterior absent; lateral oviducts placed in posterior part, part slightly upraised; posterior margin of pro- removed from each other, spermathecal gland notum slightly concave; pronotum ca. 1.4× as attached near anterior margin (as in fig. 16G); wide as long and ca. 1.9× as wide as head; meso- dorsal labiate plate without distinct tubercles, scutum slightly exposed. Hemelytron: Area along without membranous lobe posteriorly (fig. 16F). inner margin of corium swollen; inner margin of Distribution: Philippines (fig. 19). cuneus straight (Namyatova et al., 2016: fig. 13F), Host plants: Unknown. outer margin of cuneus twice as long as base. Remarks: Antennal segments and legs except Genitalia: Genital capsule (fig. 14X) twice as long all coxae and middle and hind femora are lost in as wide; ventral wall ca. 1.5× as long as dorsal females. Abdomen of males was dissected before wall, with posterior margin acute, apex placed description. almost medially, slightly curved dorsally, without Etymology: The species is named after toothlike outgrowth; left side of genital capsule Luzon Island, where the holotype was collected. not widened, margins of paramere sockets dis- Discussion: Felisacus luzonus is similar to F. tinctly angulate; distance between paramere philippinensis externally (cf. fig. 5 with 6), but the sockets ca. 0.25× as long as genital capsule width latter species differs in the markings along the at base. Right paramere (fig. 12U) distinctly inner margin of the corium where it is short, and curved; apex concave posteriorly, medial part does not extend anteriorly nor posteriorly (fig. almost as wide as basal part, without setae, with 6), the labium only slightly surpasses the poste- straight outer margin and curved inner margin rior margin of the mesosternum, the genital cap- bearing triangular outgrowth; outer angle dis- sule is broad on the left-hand side (fig. 15G), and tinct; inner angle distinct, without swellings or the ductus seminis is hooked apically (fig. 10K). outgrowth(s), bearing setae; basal part of param- Material examined: Holotype: PHILIP- ere slightly shorter than rest of paramere. Left PINES: Luzon: Dalton Pass, Nueva Vizcaya paramere (fig. 12V) irregularly shaped; apical Prov., 16.11722°N 120.96°E, 915 m, 09 Apr part flattened, with subrectangular outgrowth, 1968–10 Apr 1968, D.E. Hardy, 1♂ (00017865) removed from apex; middle side widened, with (BPBM). Paratype: PHILIPPINES: Luzon: Imu- two outgrowths, one of them bearing setae. gan [Imugin], Nueva Vizcaya [N. Viscaya] Prov., Aedeagus (general view as in Namyatova et al., 16.1589°N 120.9036°E, 949 m, 1700, Baker, 1♀ 2016: fig. 22M) conjunctiva membranous; vesica (00017867) (MZH). 114 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
A
F. magnificus F. minutus F. philippinensis F. ponaponensis
B
F. nigrescens F. ribricuneus F. schuhi F. senaru
C
F. signis F. solomonicus F. tanna F. usingeri
FIGURE 20. Distribution of Felisacus species. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 115
A
F. madagascariensis F. nigrescens F. vitilevu F. zuparkoi
B
F. wangae F. webbi F. yasunagai
C
F. bradi
FIGURE 21. Distribution of F. bradi. 116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
clade 2 clade 3 clade 14
FIGURE 22. Distribution of clades 2, 3, and 14. tudinal sulcus on dorsal surface of head pale brown; buccula, mandibular and maxillary Felisacus madagascariensis Poppius plates yellow; clypeus pale brown, brown api- Figures 5, 21 cally; tubercle around antennal fossa yellow Felisacus madagascariensis Poppius, 1912: 182 with red markings. Eyes white. Labium: Whitish (original description). yellow, segment IV slightly darker than other segments. Antenna: Segments I–II reddish Diagnosis: Recognized by the following brown. Thorax: Anterior part of pronotum combination of characters: marking along inner brown anteriorly, and yellow with reddish tinge margin of corium yellow, not reaching R+M posteriorly, posterior part of pronotum dark anteriorly and posteriorly (fig. 5); transverse brown with yellow longitudinal stripe medially; depression delimiting occipital region present mesoscutum pale brown, scutellum yellow with only dorsally; antennal segment I cylindrical (as anterior angles pale brown; thoracic pleura and in Namyatova et al., 2016: fig. 8A), ca. 1.7× as scent gland evaporative area yellow. Hemelytron: long as head width, reddish brown; pronotum Mostly opaque; inner part of clavus pale brown mostly brown to dark brown with yellow longi- with reddish tinge, outer part of clavus whitish tudinal stripe medially on posterior part; labium yellow, pale brown apically; corium whitish yel- reaching posterior margin of mesosternum; dor- low, marking along inner margin of corium yel- sal surface of labial segment II elongate posteri- low, short and narrow, not reaching R+M; orly (as in Namyatova et al., 2016: fig. 6D). embolium whitish yellow, with reddish margins; Redescription: Male. Unknown. Female. cuneus whitish yellow, with reddish outer and Total length 4.0. COLORATION (fig. 5): Head: posterior margins; membrane translucent, pale Pale brown dorsally and anteriorly with reddish brown with pale brown tinge. Legs: Coxa whit- tinge, lateral and ventral surfaces yellow; longi- ish yellow. SURFACE AND VESTITURE: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 117
Corium smooth, with very shallow punctures. Poppius (1912) described Felisacus madagas- STRUCTURE AND MEASUREMENTS: Body cariensis from a single specimen, and he noted 4.2× as long as pronotum width. Head: Depres- that it was a male. We examined the type species, sion delimiting occipital region present only housed in the MZH, which is a female, suggest- dorsally; distance from depression to pronotum ing that Poppius’ recording on the sex of the distinctly shorter than eye diameter; longitudi- specimen was incorrect. Felisacus madagas- nal sulcus on dorsal surface longer than eye cariensis is similar to F. vitilevu and F. gressitti in diameter; distance from eye to pronotum coloration and structure (figs. 5, 7). Felisacus viti- slightly longer than eye diameter, not swollen levu can be separated from F. madagascarensis by laterally; vertex ca. 2.1× as wide as eye, upraised the labium reaching the posterior margin of the (as in Namyatova et al., 2016: fig. 6D). Labium: metasternum, the anterior part of the pronotum Reaching posterior margin of mesosternum; is yellow, antennal segment I is mostly pale segments I and II strongly reduced, combined brown or reddish, ca. 1.3–1.6× as long as the length subequal to half of segment III; segment head width. Felisacus gressitti differs fromF. I shorter than wide; segment II slightly longer madagascariensis in the anterior part of the pro- than width, its dorsal surface elongate posteri- notum is yellow and the labium reaches abdomi- orly; segment IV ca. 1.5× as long as segment III. nal segment II. Antenna: Segment I cylindrical (as in Namya- Material examined: Holotype: MADAGAS- tova et al., 2016: fig. 8A), 1.7× as long as head CAR: [Madagascar], 1700, A.E. Hildebrand, 1♀ width, 1.0× as long as pronotum width. Thorax: (00018976) (MZH). Anterior part shorter than posterior part; collar not delimited; posterior part slightly upraised; Felisacus magnificus Distant, 1904 posterior margin concave, pronotum ca. 1.4× as wide as long and ca. 1.7× as wide as head; meso- Figures 6, 10J, 12W, X, 14Y, 16A, B, 20 scutum exposed. Hemelytron: Area along inner Felisacus magnificus Distant, 1904: 439 (original margin of corium almost flat; inner margin of description). corium convex (as in Namyatova et al., 2016: fig. Felisacus pulchellus Poppius, 1915a: 80 (synony- 13E), outer margin of corium almost ca. 3× as mized by Carvalho, 1981: 62). long as base. Genitalia: Dorsal labiate plate Felisacus capitatus Miyamoto, 1965: 164; new wider than distance between apodemes of sec- synonymy. ond valvulae; with semicircular sclerite and dis- tinct sclerotized rings laterally (similar to Diagnosis: Recognized by the following Namyatova et al., 2016: fig. 23F); dorsal labiate combination of characters: presence of distinct plate with distinct tubercles (similar to Namya- C-shaped dark brown marking on corium, with tova et al., 2016: fig. 23G). anterior and posterior parts reaching R+M, ante- Distribution: Madagascar (fig. 21). rior part not inclined posteriorly (fig. 6); anten- Host plants: Unknown. nal segment I widened (as in Namyatova and Discussion: Antennal segments III–IV, Cassis, in press: fig. 8B), red; antennal segment II femora, tibiae, and tarsi are lost. The abdomen often brown; dorsal surface of head flat (as in is in a vial attached to the specimen. Vestiture Namyatova et al., 2016: fig. 6E), vertex flat; of the holotype has been lost. Some characters labium reaching posterior margin of mesoster- of the female genitalia were not recorded, num; apex of cuneus red; femora often with including striation of the dorsal labiate plate, brown or red markings in apical half; ventral position of the lateral oviducts, spermathecal wall of genital capsule with tooth posteriorly (fig. gland, and presence of the membranous lobe 14Y); right paramere straight, its inner angle of the posterior wall. with slightly curved outgrowth (fig. 12W); apical 118 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 part of left paramere widened, with tooth api- very pale; tibiae often yellow basally and whitish cally and without outgrowth on dorsal surface; yellow apically, sometimes pale brown, brown, medial part of left paramere with narrow out - or reddish basally and yellow apically, rarely growth (fig. 12X); apical half of ductus seminis uniformly yellow; foretarsus whitish or yellow, sclerotized, hooked apically; vesica without scler- middle and hind tarsi yellow. Abdomen: Yellow ites (fig. 10J). with brown segment IX and dorsum, sometimes Redescription: Male. Total length 3.4–4.0. reddish apically. SURFACE AND VESTITURE: COLORATION (fig. 6): Head: Mostly yellow to Corium smooth, with shallow punctation. Dor- pale brown, sometimes with reddish markings; sum clothed with setae longer than antennal buccula and apex of clypeus often brown. Eye segment II diameter; antennal segment I clothed dark brown with reddish tinge, sometimes uni- with suberect setae shorter than antennal seg- formly red. Labium: Segment I–II brown, seg- ment II diameter; femora clothed with suberect ments III yellow to pale brown, rarely brown, setae subequal to or longer than antennal seg- segment IV yellow. Antenna: Segment I red, red- ment II diameter; abdomen clothed with suber- dish brown or brown, segment II brown, often ect setae of different length. STRUCTURE AND yellow at base, rarely yellow and brown apically, MEASUREMENTS: Body ca. 3.9–4.2× as long segments III–IV pale brown to brown, often as pronotum width. Head: Transverse depres- darker than segment II, sometimes segment IV sion delimiting occipital region present only somewhat paler than segment III. Thorax: Ante- dorsally; distance between depression and pro- rior part of pronotum yellow to pale brown, notum slightly longer than eye diameter; longi- brown near forecoxae, posterior part of prono- tudinal sulcus on dorsal surface of head shorter tum uniformly brown to dark brown; scutellum than eye diameter; eye diameter distinctly and mesoscutum brown to dark brown; thoracic shorter than distance from head to pronotum; pleura mostly brown to dark brown, metapleu- vertex ca. 1.9–2.3× as wide as eye, flat (Namya- ron often whitish yellow posteriorly, scent gland tova et al., 2016: fig. 6E). Labium: Reaching pos- evaporative area whitish yellow. Hemelytron: terior margin of mesosternum or slightly Mostly translucent; clavus opaque, uniformly surpassing it; segments I and II as long as wide, brown to dark brown; corium, embolium, and combined ca. 1.5× as long as segment III; dorsal cuneus translucent; corium with yellow tinge, surface of segment II not elongate posteriorly with brown anterior angles and C-shaped dark (as in Namyatova et al., 2016: fig. 6E); segment brown marking, anterior part of this marking III shorter than ventral side of head; segment IV reaching R+M, posterior part reaching costal subequal to half of segment III. Antenna: Seg- margin; embolium colorless, with dark brown ment I widened medially (as in Namyatova et apex and sometimes pale brown to dark brown al., 2016: fig. 8B), ca. 1.1–1.2× as long as head marking medially, margins of embolium yellow width, ca. 0.6–0.7× as long as pronotum width; to pale brown, outer margin sometimes some- segment II ca. 1.4–1.6× as long as head width, what darker than inner margin; cuneus colorless ca. 0.7–0.8× as long as pronotum width; seg- with red apex, yellow outer margin and colorless ment III slightly longer than segment II and ca. inner margin; membrane gradually changing twice as long as segment IV. Thorax: Anterior color from pale brown to brown anteriorly and part of pronotum distinctly shorter than poste- yellow posteriorly, membrane cell pale brown to rior part; collar distinct; posterior part slightly brown. Legs: Coxae whitish yellow to yellow; upraised; posterior margin of pronotum usually femora yellow, often whitish yellow basally, often straight, sometimes slightly concave; pronotum hind femur and rarely fore- and middle femora ca. 1.2–1.4× as wide as long and ca. 1.8–2.9× as with pale brown to brown or reddish apex and wide as head; mesoscutum usually slightly marking on apical part, sometimes that marking exposed, sometimes not exposed. Hemelytron: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 119
Area along inner margin of corium; inner mar- gin; dorsal labiate plate without distinct tuber- gin of cuneus straight (as in Namyatova et al., cles, without membranous lobe posteriorly. 2016: fig. 13F), outer margin of cuneus twice as Distribution: Japan, Java, Taiwan, Vietnam, long as base. Abdomen: Genital capsule rotated (fig. 20). 45º. Genitalia: Genital capsule (fig. 14Y) twice as Host plants: Some specimens from Lombok long as wide; ventral wall ca. 1.5× as long as dor- Is. were collected from Plesioneuron sp. (Thelyp- sal wall, with posterior margin of ventral wall teridaceae) and Diplazium esculentum nor curved, with toothlike outgrowth; right side (Woodsiaceae). of genital capsule folded, left side of genital cap- Discussion: Distant (1904) described F. sule not widened; paramere sockets angulate; magnificus from Burma. Poppius (1915a) distance between paramere sockets ca. 0.3× as described F. pulchellus from the Philippines, long as genital capsule width at base. Right para- noting that it differs from F. magnificus by its mere (fig. 12W) almost straight; apex slightly different coloration, antennal structure, and concave; medial part narrower than basal part, smooth basal part of the pronotum. Poppius without setae, outer margin convex and inner (1915b) described F. longiceps from Taiwan and margin concave, without swelling; outer angle compared it with F. pulchellus. According to absent; inner angle distinct with curved out- him, F. longiceps differs from the latter by the growth, bearing setae; basal part ca. 0.7× as long longer head and antennal segments, and differ- as rest of paramere. Left paramere (fig. 12X) dis- ent leg coloration. Carvalho (1981) synony- tinctly curved medially; apical part flattened, mized Felisacus pulchellus with F. magnificus. with toothlike outgrowth apically, posterior sur- We examined the holotypes of F. longiceps face without other outgrowth; middle part wid- (female), and F. pulchellus (female) and lecto- ened dorsally, with narrow outgrowth at inner type of F. magnificus (sex unknown) and regard side, bearing setae. Aedeagus (general view as in them as very similar to each other. As all the Namyatova et al.: fig. 22M) conjunctiva mem- types are females or had lost genitalia, it is branous; vesica without spicules; apical half of problematic to determine which specimens ductus seminis sclerotized, secondary gonopore belong to these three species. We also examined placed near phallotheca mouth in repose; ductus many specimens similar to these species from seminis hooked apically (fig. 10J). Southeast Asia. We were able to split the speci- Female. Total length 3.6–4.0. COLORATION, mens into three species, based on male genita- SURFACE, AND VESTITURE (fig. 6): As in lia. They also slightly differ in coloration. The male. STRUCTURE AND MEASUREMENTS: larger specimens with the dark brown to black Structure as in male; body ca. 4.0–4.2× as long as antennal segment II and bands on the hind pronotum width; vertex ca. 1.9–2.1× as wide as femora most likely belong to F. magnificus, and eye; antennal segment I ca. 1.1–1.2× as long as are very similar to the type specimen, preserved head width, ca. 0.6× as long as pronotum width; in BMNH, and to the type of its synonym F. segment II ca. 1.5–1.7× as long as head width, ca. pulchellus, preserved in the HNHM. The rest of 0.8–0.9× as long as pronotum width; pronotum the specimens with yellow antennal segment II ca. 1.2–1.4× as wide as long and ca. 1.9–2.1× as and the femora without banding represent two wide as head. Genitalia (figs. 16A, B): Dorsal species, with one of them having an identical labiate plate very small and transparent, as wide vesica compared with F. magnificus, and the as distance with apodemes of second valvulae, other species has a unique claw-shaped spicule, without striations or membranous ridge medi- which has not been previously found in any ally; semicircular sclerite and sclerotized rings Felisacus species. Felisacus longiceps is without absent; lateral oviducts placed in anterior part; claw-shaped spicules and is also broadly dis- spermathecal gland attached near anterior mar- tributed in Southeast Asia. Specimens with 120 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 claw-shaped spicules were collected from the Diplazium esculentum Swartz (Woodsiaceae), Philippines and Borneo only. We regard the Tai- det. Michael Lovave (LAE herbarium, PNG), wanese specimens as conspecific and we treat 1♂ (00386604) (ZISP). Lombok Island, Senaru, the species with a yellow antennal segment II track to Tiu Kelep Waterfall, 8.30111°S and without bands on hind femora as F. longi- 116.40833°E, 31 Aug 2012, F. Konstantinov, Ple- ceps, and the species with similar color pattern sioneuron sp. (Thelypteridaceae), det. Michael but with claw-shaped spicule as the new species Lovave (LAE herbarium, PNG), 3♂ (00386636– F. zuparkoi (see below). 00386638) (ZISP). Lombok Island, nr Senaru, Felisacus longiceps is also very similar to F. capi- 8.31956°S 116.405°E, 31 Aug 2012, F. Konstan- tatus externally and the male genitalia are depicted tinov, Diplazium esculentum Swartz (Woodsia- by Miyamoto (1965: figs. 39–41), and they are ceae), det. Michael Lovave (LAE herbarium, likely conspecific. According to this author, the PNG), 3♂ (00386623, 00386624, 00386622), 2 types were preserved in the Entomological Labo- sex unknown (00386625, 00271349) (ZISP). ratory, Kyushu University, but we could not get JAPAN: Okinawa: Ishigaki Is., Ryukyu Islands, any information on them from this university. 24.4°N 124.2°E, 13 Oct 1999, Belokobylskij, 1♂ Although, according to Miyamoto’s (1965) (00271512) (ZISP); 14 Oct 1999, Belokobylskij, description of F. capitatus, antennal segment II is 1♂ (00271511) (ZISP); 20 Oct 1999, Belokob- yellow, we consider this difference as minor and ylskij, 1♂ (00271510) (ZISP); 21 Oct 1999, synonymize F. capitatus with F. magnificus. Belokobylskij, 1♂ (00271513), 1♀ (00271506) Based on the structure of the parameres as (ZISP). TAIWAN: Pihu, 49 km E of Taipei on depicted in Carvalho (1981), the specimens he Hwy 9 to Yilan, 24.79194°N 121.78166°E, 380 examined for the description of F. magnificus are m, 18 Feb 1972, T.C. Maa, 3♂ (00043195, most likely F. longiceps. However, the external 00043197, 00043198), 7♀ (00043199–00043202, view of this species is most similar to F. magnifi- 00043204–00043206) (BPBM). Wulai nr Taipei, cus. It is also possible that Carvalho had repre- 400 m, 12 Apr 1960, T.C. Maa, 1♂ (00043196) sentatives of both species in his series, as those (BPBM). THAILAND: Nakhon Nayok:Sarika species can be collected together (see Material Waterfall, 14.30489°N 101.25506°E, 17 Jun Examined sections) The identification of F. mag- 2009–18 Jun 2009, T. Yasunaga, 5♂ (00021594– nificus in Woodward (1954) is correct. 00021598), 1♀ (00021599) (TYCN). VIET- Felisacus magnificus is also similar to F. cey- NAM: Dak Lak: Buon Ma Thuot [BanMeThuot], lonicus and F. lindbergae, and comparisons are 12.6733°N 108.04459°E, 500 m, 20 Dec 1960–24 provided in the discussion sections for those Dec 1960, C.M. Yoshimoto, 1♂ (00043193) two species. (BPBM). Gia Lai: 20 km N Kan Nak [Zalay- Material examined: Lectotype (Felisacus Kontum Prov., Buon-Luoi Dist., 20 km N Kan- magnificus): MYANMAR: Myitta, Tenasserim nack], 14.18252°N 108.58748°E, 20 Oct 1988, Valley, 14.1667°N 98.5167°E, 183 m, Doherty, Sharkov, 1♀ (00271526) (ZISP). Hòa Bình: Lectotype, 1 sex unknown (BMNH). Holotype Hoa Binh: Near Mai Chau [Mai Chon, Mai (F. pulchellus): Laguna: Los Banos, 1700, Baker, Chou], 20.667°N 105.084°E, 31 Oct 1990, Belo- 1♀ (00018445) (MZH). Additional material: kobylskij, 4♂ (00271522–00271525) (ZISP). INDONESIA: Java: Bogor, 6.58916°S Lam Dong: Fyan [Ngoc Son], 11.88333°N 106.79305°E, 240 m, 23 Apr 1955, J. v. d. Vecht, 108.2°E, 1200 m, 11 Jul 1961–09 Aug 1961, N.R. 2♂ (00018971, 00018968), 1 sex unknown Spencer, 1♂ (00043194) (BPBM). Vinh Phuc: (00018970), 1♀ (00018969) (NML).West Nusa Tam Dao, 100 km NW Ha Noi [Hanoi], Tenggara: Lombok Island, Senaru, Panorama 21.47607°N 105.56339°E, 1000 m, 16 Feb 1990, track to Sendang Gila Waterfall, 8.30111°S Belokobylskij, 4♀ (00045813, 00045812, 116.40833°E, 30 Aug 2012, F. Konstantinov, 00045811, 00045810), 6♂ (00045814, 00045812, 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 121
00045810) (ZISP); 12 Nov 1990, Belokobylskij, opaque, red; membrane colorless with pale 1♀ (00045815), 1♂ (00045815) (ZISP); 14 Nov brown cell. Legs: Mostly yellow, with tarsal seg- 1990, Belokobylskij, 11♂ (00038596–00038598, ments II and III often yellow to pale brown. 00386602, 00386601, 00038599), 6♀ (00038597, Abdomen: Whitish yellow to yellow with brown 00038598, 00386602, 00038599) (ZISP). dorsal surface and segment IX. SURFACE AND VESTITURE: Corium smooth, with shallow and scarce punctures. Dorsum with suberect setae Felisacus malayensis, sp. nov. subequal to or shorter than antennal segment II Figures 6, 9A, 12Y, Z, 15A, 19 diameter; antennal segment I and femora with Diagnosis: Recognized by the following suberect setae shorter than antennal segment II combination of characters: head and pronotum diameter; abdomen clothed with suberect, mostly yellow to pale brown with posterior part mostly short simple setae. STRUCTURE AND of pronotum mostly brown, cuneus red; mark- MEASUREMENTS: Body 4.0–4.4× as long as ing along inner margin of corium brown, not pronotum width. Head: Depression delimiting reaching R+M anteriorly; cylindrical antennal occipital region present dorsally and laterally (as segment I (as in Namyatova et al., 2016: fig. 8A), in Namyatova et al., 2016: fig. 4E); distance transverse depression on head extending later- between depression and pronotum distinctly ally, vertex upraised (as in Namyatova et al., shorter than eye diameter; longitudinal sulcus on 2016: fig. 6D), body length in male 3.2–3.7, in dorsal surface longer than eye diameter distance female 3.8; labium reaching posterior margin of from eye to pronotum eye slightly longer than metasternum or slightly surpassing it; cuneus ca. eye diameter, not swollen laterally (as in Namya- 3× as long as base; medial part of right paramere tova et al., 2016: fig. 4E); vertex ca. 1.6–1.9× as only slightly wider than basal part, shorter than wide as eye; upraised (as in Namyatova et al., basal and apical parts combined (fig. 12Y); 2016: fig. 6D); buccula ca. 0.2–0.25× as long as vesica with five spicules, including spicules A, B, clypeus. Labium: Reaching posterior margin of and C (fig. 9A). mesosternum or slightly surpassing it; segments Description: Male. Total length 3.2–3.7. I and II strongly reduced, combined shorter than COLORATION (fig. 6): Head: Mainly pale half of segment III; segment I shorter than wide brown, anterior part of head and frons with red- (as in Namyatova et al., 2016: figs. 6D, 9C); seg- dish tinge. Eye dark brown, reddish at sides. ment II longer than wide, its dorsal surface elon- Labium: Uniformly yellow. Antenna: Segment I gate posteriorly; segment III slightly longer than yellow, darker apically; segment II pale brown, ventral side of head; segment IV ca. 1.5× as long brown apically; segments III–IV brown to dark as segment III. Antenna: Segment I cylindrical brown. Thorax: Anterior part of pronotum pale (as in Namyatova et al., 2016: fig. 8A), ca. 1.6– brown with collar brown; posterior part of pro- 1.7× as long as head width, ca. 1.1× as long as notum mostly brown, pale brown anteriorly; pronotum width; segment II ca. 1.8–2.0× as long mesoscutum and scutellum pale brown, scutel- as head width, ca. 1.2–1.3× as long as pronotum lum brown apically; thoracic pleura brown; scent width; segment III slightly longer than segment gland evaporative area yellow, pale brown api- II; segment IV ca. 0.4–0.5× as long as segment cally. Hemelytron: Mostly translucent; inner part IV. Thorax: Anterior part of pronotum slightly of clavus opaque, pale brown; outer part of cla- shorter than posterior part; collar delimited; pos- vus colorless, yellow to pale brown apically; terior part slightly upraised; posterior margin of corium colorless, whitish with marking along pronotum slightly concave; pronotum ca. 1.2– inner margin of corium brown; embolium mostly 1.3× as wide as long and ca. 1.5× as wide as head; colorless, with yellow apical part and reddish mesoscutum exposed. Hemelytron: Area along apex, margins pale brown; cuneus sometimes inner margin of corium almost flat; inner margin 122 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 of cuneus convex (as in Namyatova et al., 2016: I ca. 1.4× as long as head width, ca. 0.9× as long fig. 13E), outer margin ca. 3× as long as base. as pronotum width; segment II ca. 1.9× as long Abdomen: Genital capsule rotated at right angle as head width, ca. 1.2× as long as pronotum relative to rest of abdomen. Genitalia: Genital width; pronotum ca. 1.3× as wide as long and ca. capsule (fig. 15A) twice as long as wide; ventral 1.6× as wide as head. Genitalia (as in Namyatova wall ca. 1.5× as long as dorsal wall, its posterior et al., 2016: fig. 23F, G): Dorsal labiate plate margin smooth, semioval, without outgrowth(s), wider than distance between apodemes of sec- not curved dorsally; its apex inclined rightward; ond valvula; mostly smooth, without distinct sides of genital capsule not modified; margins of striations, with semicircular sclerite and distinct paramere sockets more or less rounded; distance sclerotized rings laterally; lateral oviducts placed between paramere sockets ca. 0.7× as long as almost medially, very close to each other, sper- genital capsule base. Right paramere (fig. 12Y) mathecal gland placed between lateral oviducts; distinctly curved in apical half; apex slightly con- dorsal labiate plate with distinct tubercles, with- cave; medial part slightly wider than basal part, out membranous lobe medially. Antennae are bearing setae, with outer margin concave and lost in females. inner margin widened; outer angle distinct, wid- Distribution: Malaya (Penang) (fig. 19). ened; inner angle rounded, without setae; basal Host plants: Unknown. part of paramere ca. 0.15–0.2× as long as rest of Etymology: The species is named after the paramere. Left paramere (fig. 12Z) L-shaped; Malaya Peninsula, where it was collected. apical part flattened, with tooth on posterior side Discussion: Felisacus malayensis is similar to medially and without outgrowth on dorsal sur- F. bau, F. bellus, and F. jacobsoni in coloration face; middle part widened, without swelling or (cf. figs. 4, 5, 6) and shape of the right paramere outgrowth; setae only on middle part near outer having a long apical process (cf. fig. 12Y with margin. Aedeagus (general view as in Namyatova figs. 11H, J, 12E). Felisacus jacobsoni differs from et al., 2016: fig. 22I) conjunctiva weakly sclero- F. malayensis in the larger size, the body length tized, secondary gonopore placed at base of is 4.4–4.7 mm in males and 4.6–4.8 mm in vesica in repose; sclerotization of ductus seminis females, and in the shape and configuration of around secondary gonopore shorter than wide; the spicules (fig. 8M). Felisacus bellus can be vesica with five spicules, including spicules A, B, separated from F. malayensis in the presence of and C (fig. 9A). six vesical spicules, including spicules A, B, and Female. Total length 3.8. COLORATION G (fig. 8E). Felisacus. bau differs from F. malay- (fig. 6): Mostly pale brown, dorsal surface, frons, ensis in the cuneus twice as long as its base and and areas below antennal fossa with reddish the presence of six vesical spicules, including tinge, antennal segment I with reddish tinge, spicules A and C (fig. 8D). antennal segment II brown to dark brown, inner Material examined: MALAYSIA: Pulau part of clavus yellow. Thorax and legs: As in Pinang [Penang], 5.26553°N 100.48428°E, 12 Jan male. Hemelytron: Similar to male, but outer 1959–14 Jan 1959, L.W. Quate, 2♂ (00017857, part of clavus pale brown, brown apically; 00017858), 1♀ (00017859) (BPBM). cuneus red with brown outer margin and paler outer part; membrane with pale brown tinge, Felisacus meilingae, sp. nov. membrane cell brown. Abdomen: Yellow with segments VIII–IX and dorsal surface red. SUR- Figures 6, 9B, 12AA, AB, 15B, 19 FACE AND VESTITURE: As in male. STRUC- Diagnosis: Recognized by the following TURE AND MEASUREMENTS: Structure as in combination of characters: dorsal surface of head male; body ca. 4.0× as long as pronotum width; and pronotum mostly yellow, lateral margins of vertex ca. 2.2× as wide as eye; antennal segment collar not dark; inner part of clavus often pale 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 123 brown to brown, sometimes yellow with brown cell, rarely cell colorless. Legs: Coxae whitish or margins; posterior margin of pronotum with yellow, femora whitish yellow to yellow basally brown marking laterally; cuneus mostly translu- and darker, yellow to pale brown or reddish api- cent, colorless (fig. 6); cylindrical antennal seg- cally; tibia yellow to pale brown, often paler api- ment I (as in Namyatova et al., 2016: fig. 8A), cally, sometimes with reddish tinge; tarsi pale transverse depression on head extending later- brown to brown, sometimes segment I yellow ally, vertex upraised, labium reaching middle of and segments II–III pale brown or brown. Abdo- mesosternum or slightly surpassing it; antennal men: Whitish yellow to yellow, often with dorsal segment I ca. 1.5–1.7× as long as wide; labium surface pale brown, brown, or red. SURFACE reaching middle of mesosternum; cuneus ca. 3× AND VESTITURE: Corium smooth, with shal- as long as base; medial part of right paramere low and scarce punctures. Dorsum, antennal seg- slightly wider than basal part, shorter than basal ment I and femora with setae subequal to or and apical parts combined (fig. 12AA); apical slightly longer than antennal segment II diame- part of left paramere not widened, ca. 3× as long ter; abdomen clothed with short erect setae, dis- as wide (fig. 12AB); vesica with six spicules, tinctly mostly shorter than antennal segment II including spicules A, C, D, G, and I (fig. 9B). diameter. STRUCTURE AND MEASURE- Description: Male. Total length 4.0–4.5. MENTS: Body ca. 4.2–4.4× as long as pronotum COLORATION (fig. 6): Head: Whitish yellow to width. Head: Depression delimiting occipital yellow, dorsal surface often somewhat darker region distinct dorsally and laterally (as in Namy- than other sides; head sometimes with stripe atova et al., 2016: fig. 4E); distance between above eye and marking between eyes reddish and depression and pronotum distinctly shorter than clypeus with darker apex. Eye dark brown to eye diameter; longitudinal sulcus on dorsal sur- black, often with reddish tinge.Labium: Yellow, face of head longer than eye diameter; distance segment III often brown or reddish ventrally. from eye to pronotum longer than eye diameter, Antenna: Segment I yellow to brown with paler not swollen laterally (as in Namyatova et al., base, segment II pale brown to brown, usually 2016: fig. 4E); vertex ca. 1.6–2.0× as wide as eye, darker than segment I, segments III–IV brown to upraised (as in Namyatova et al., 2016: fig. 6D). dark brown. Thorax: Pronotum mostly yellow, Labium (as in Namyatova et al., 2016: figs. 6D, with anterior margin pale brown or brown and 9C): Reaching middle of mesosternum; segments posterior margin with pair brown to dark brown I and II strongly reduced, combined shorter than marking near humeral angle, sometimes pale half of segment III; segment I shorter than wide; brown between markings; scutellum and mesos- segment II as long as wide, its dorsal surface cutum whitish yellow to pale brown, scutellum elongate posteriorly; segment III longer than with brown apex; thoracic pleura yellow to pale ventral side of head; segment IV ca. 1.5× as long brown, scent gland evaporative area whitish yel- as segment III. Antenna: Segment I cylindrical low to yellow, often darker apically, paler than (as in Namyatova et al., 2016: fig. 8A), ca. 1.5– thoracic pleura. Hemelytron: Mostly colorless and 1.7× as long as head width, ca. 0.9–1.2× as long translucent; inner part of clavus yellow to brown as pronotum width; segment II ca. 1.6–2.5× as with small markings and margins pale brown to long as head width, ca. 1.1–1.5× as long as pro- brown; outer part of clavus sometimes with pale notum width; segment III slightly longer than brown to brown apex; corium with marking long, segment II; segment IV ca. 0.3× as long as seg- inner margin of corium yellow to dark brown, ment III. Thorax: Anterior and posterior parts of often paler medially; embolium sometimes yel- pronotum subequal in length, collar delimited; low with brownish margins; cuneus with brown posterior part slightly upraised; posterior margin outer and posterior margins and whitish inner of pronotum concave, pronotum ca. 1.1–1.4× as margin; membrane with yellowish to pale brown wide as long and ca. 1.4–1.8× as wide as head; 124 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 mesoscutum exposed. Hemelytron: Area along male. STRUCTURE AND MEASUREMENTS: inner margin of corium almost flat; inner margin Structure as in male; body ca. 4.2–4.4× as long as of cuneus convex (as in Namyatova et al., 2016: pronotum width; vertex ca. 1.6–2.0× as wide as fig. 13E), outer margin of cuneus ca. 3× as long eye; antennal segment I ca. 1.4–1.7× as long as as base. Abdomen: Genital capsule rotated left- head width, ca. 0.8–1.1× as long as pronotum ward at right angle relative to rest of abdomen. width; segment II ca. 1.8–2.1× as long as head Genitalia: Genital capsule (fig. 15B) ca. 1.5× as width, ca. 1.1–1.3× as long as pronotum width; long as wide; ventral wall ca. 1.5× as long as dor- pronotum ca. 1.2–1.3× as wide as long and ca. sal wall, with posterior margin of ventral wall 1.5–1.7× as wide as head. Genitalia (as in Namya- semioval, smooth, without outgrowth(s), its apex tova et al., 2016: fig. 23F, G): Dorsal labiate plate inclined to left-hand side, not curved; left side of wider than distance between apodemes of second genital capsule not widened; right paramere valvula; mostly smooth, without distinct stria- socket slightly acute, left one rounded; distance tions, with semicircular sclerite and distinct scler- between paramere sockets subequal to half of otized rings laterally; lateral oviducts placed genital capsule width at base. Right paramere almost medially, very close to each other, sperma- (fig. 12AA) distinctly curved in apical half; apex thecal gland placed between lateral oviducts; dor- slightly concave; medial part slightly wider than sal labiate plate with distinct tubercles, without basal part, bearing setae, with outer margin membranous lobe medially. slightly concave and inner margin convex; outer Distribution: Vanuatu (Ambrym Is., Pente- angle distinct; inner angle indistinct, not bearing cost Is., Shepherd Is., and Maewo Is.) (fig. 19). setae; basal part of paramere ca. 0.15–0.2× as Host plants: Unknown. long as rest of paramere. Left paramere (fig. Etymology: This species is named after Mei- 12AB) L-shaped; apical part not flattened, with Ling Chan (National Museum of Natural Sci- tooth on posterior side medially (as in fig. 11G) ences, Taichung, Taiwan), who loaned us the and without outgrowth on dorsal surface; middle paratype of Felisacus bellus Lin, 2000. part widened, without swelling or outgrowth; Discussion: Felisacus meilingae is similar to setae only on middle part near outer margin. F. schuhi in coloration and structure (fig. 6), but Aedeagus (general view as in Namyatova et al., the latter species can be separated by the prono- 2016: fig. 22I) conjunctiva weakly sclerotized; tum uniformly yellow to pale brown and the secondary gonopore placed at base of vesica in presence of five vesical spicules (fig. 9J). repose; sclerotization of ductus seminis around Felisacus meilingae is also similar to F. myersi in secondary gonopore shorter than wide; vesica external morphology (fig. 6) and the shape of vesi- with six spicules, including spicules A, C, D, G, I cal spicules (cf. fig. 9B with 9C); F. myersi differs (fig. 9B). from it by the dorsal surface of the head brown to Female. Total length 4.2–4.6. COLORATION dark brown, the inner part of the clavus mostly col- (fig. 6): Head: Similar to male, but coloration orless or yellow and it has four vesical spicules. varying from whitish yellow to pale brown. Material examined: Holotype: VANUATU: Labium: As in male. Antenna: Segments I–II often Malampa: Ambrym Is.: Ranon to Mount Toyo, as in males, but rarely reddish. Thorax: Similar to 16.15949°S 168.13365°E, 400 m, 02 Sep 1979, male, but posterior part of pronotum, mesoscu- W.C. Gagne, 1♂ (00043085) (BPBM). Paratypes: tum, and scutellum sometimes with reddish tinge; VANUATU: Malampa: Ambrym Is.: Ambrym thoracic pleura yellow to pale brown. Hemelytron: Is., 16.24546°S 168.12298°E, Dec 1984, N.L.H. Similar to male, but inner part of clavus some- Krauss, 1♂ (00043095) (BPBM). Ranon to times uniformly brown. Legs: As in male. Abdo- Mount Toyo, 16.15949°S 168.13365°E, 400 m, 02 men: Yellow to pale brown with reddish dorsal Sep 1979, W.C. Gagne, 5♂ (00043084, surface. SURFACE AND VESTITURE: As in 0004308611H, J, 00043089), 1♀ (00043090), 1 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 125 sex unknown (00043091) (BPBM). S Ambrym with wide yellow stripe medially; mesoscutum [Ambrim] Is., 16.35815°S 168.14838°E, 22 Aug and scutellum yellow; thoracic pleura yellow; 1967–04 Sep 1967, J. and M. Sedlacek, 1♀ scent gland evaporative area yellow with reddish (00043092) (BPBM). Penama: Maewo Is.: Soun- tinge apically. Hemelytron: Mostly translucent; wari, 15.38°S 168.12989°E, 180 m, 04 Sep 1979– inner part of clavus opaque, yellow with reddish 05 Sep 1979, G.A. Samuelson, 2♀ (00043064, tinge and with red margins; corium colorless; 00043065) (BPBM). Sounwari, 15.38°S marking along inner margin of corium pale 168.12989°E, 20 m, 04 Sep 1979, B.H. Gagne, 4♂ brown; cuneus red, with yellow outer part and (00043048–00043051), 6♀ (000430055– brown margins; membrane and membrane cell 00043060), 1 sex unknown (00045806) (BPBM). with grayish tinge. Legs: Mostly yellow with red Pentecost Is.: Wall Bay, 15.76364°S 168.19233°E, markings in femora and tibiae; tarsal segments 50 m, Dec 1984, N.L.H. Krauss, 1♀ (00043096) II–III pale brown. SURFACE AND VESTITURE: (BPBM). Shefa: Shephard Group: Tongariki Corium with shallow scarce punctures. Setae on Island, 16.8°S 168.5°E, 300 m, 29 Aug 1979, G.M. body shorter or as long as antennal segment II Nishida, 4♀ (00042274, 00044635, 00043094, diameter. STRUCTURE AND MEASURE- 00043093) (BPBM). MENTS: Body ca. 4.3× as long as pronotum width. Head: Depression delimiting occipital region distinct dorsally and laterally (as in Felisacus minutus Carvalho Namyatova et al., 2016: fig. 4E); distance between Figures 6, 9D, 12AC, AD, 20 depression and pronotum distinctly shorter than Felisacus minutus Carvalho, 1981: 6 (original eye diameter; longitudinal sulcus on dorsal sur- description). face of head longer than eye diameter; distance from eye to pronotum subequal to eye diameter, Diagnosis: Recognized by the following com- not swollen laterally (as in Namyatova et al., bination of characters: short body, 2.6 mm in 2016: fig. 4E); vertex ca. 1.3× as wide as eye, male; posterior margin of pronotum brown with upraised (as in Namyatova et al., 2016: fig. 6D). yellow wide stripe medially; cuneus red, with yel- Labium (as in Namyatova et al., 2016: figs. 6D, low outer part and brown margins; marking along 9C): Segments I and II strongly reduced, com- inner margin of corium pale brown, not reaching bined length shorter than half of segment III; R+M anteriorly antennal segment I cylindrical (as segment I shorter than wide; segment II slightly in Namyatova et al., 2016: fig. 8A); transverse longer than wide, its dorsal surface elongate pos- depression on head extending laterally, vertex teriorly; segment III subequal to ventral side of upraised (as in Namyatova et al., 2016: fig. 6D); head. Antenna: Segment I cylindrical (as in vertex ca. 1.3× as wide as eye diameter; cuneus 3× Namyatova et al., 2016: fig. 8A), ca. 1.6× as long as wide as base; medial part of right paramere dis- as head width, ca. 1.1× as long as pronotum tinctly wider than basal part, its outer and inner width; segment II ca. 2.0× as long as head width, parts straight (fig. 12AC); vesica with six spicules, ca. 1.4× as long as pronotum width. Thorax: including spicule A. Anterior part of pronotum slightly shorter than Redescription: Male. Total length 2.6. COL- posterior part; collar delimited; posterior part ORATION (fig. 6): Head: Pale brown with red- slightly upraised; posterior margin of pronotum dish tinge. Eye brown, reddish laterally. Labium: concave; pronotum ca. 1.2× as wide as long and Yellow. Antenna: Segment I pale brown with red- ca. 1.4× as wide as head; mesoscutum exposed. dish tinge, whitish yellow basally; segment II Hemelytron: Area along inner margin of corium reddish brown. Thorax: Anterior part of prono- flat; inner margin of cuneus convex (as in Namy- tum yellow with reddish tinge and brown ante- atova et al., 2016: fig. 13E), outer margin of rior margin; posterior part of pronotum brown cuneus ca. 3× as long as base. Genitalia: Right 126 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 paramere (fig. 12AC) distinctly curved in apical p ale brown to brown margins; cylindrical part; apex straight dorsally; medial part distinctly antennal segment I (as in Namyatova et al., wider than basal part, bearing setae, with mar- 2016: fig. 8A), transverse depression on head gins almost straight; outer angle distinct, inner extending laterally, vertex upraised (as in angle rounded, without setae; basal part of para- Namyatova et al., 2016: fig. 6D). setae on fem- mere ca. 0.3× as long as rest of paramere. Left ora mostly longer than antennal segment II paramere (fig. 12AD) L-shaped; apical part not width; labium reaching middle of mesosternum flattened, with tooth on posterior side medially or slightly surpassing it; cuneus ca. 3× as long (as in fig. 34G) and without outgrowth on dorsal as base; medial part of right paramere slightly surface; middle part widened, without swelling wider than basal part, shorter than basal and or outgrowth; setae only on middle part near apical parts combined (fig. 12AE); apical part of outer margin. Aedeagus (general view as in left paramere not widened, ca. 3× as long as Namyatova and Cassis, in press: fig. 22I) con- wide (fig. 12AF); vesica with five spicules, junctiva weakly sclerotized, sclerite around sec- including spicules A, C, D, G, I (fig. 9C). ondary gonopore short, bowl shaped, secondary Description: Male. Total length 3.4–4.2. gonopore placed at base of vesica in repose; COLORATION (fig. 6): Head: Dorsal surface of vesica with six spicules, including long spicule A head brown to dark brown, yellow posteriorly, and five additional spicules of different shape anterior side of head pale brown to brown, some- and size (fig. 9D). times with yellow markings below inferior mar- Female. Not seen. gin of antennal fossa; clypeus pale brown to dark Distribution: Papua New Guinea (fig. 20). brown, darkened apically; tubercles around Host plants: Unknown. antennal fossa yellow to pale brown, sometimes Discussion: Carvalho (1981) described F. brown dorsally; mandibular plate often yellow to minutus from two males. We examined the holo- brown; maxillary plate whitish yellow to pale type and an additional male specimen of the spe- brown; buccula whitish yellow to yellow; ventral cies. This species is unlike its congeners on the side of head yellow. Eye whitish yellow to brown, basis of coloration of the pronotum, size of the sometimes with reddish tinge. Labium: Often vesical spicules. uniformly whitish yellow to yellow, sometimes Material examined: Holotype: PAPUA segment I yellow to pale brown, rarely segment NEW GUINEA: Manus: Bismarck Arch., II brownish basally or labium uniformly pale Momote, 2.06168°S 147.4245°E, 24 Dec 1959, brown. Antenna: Segment I often pale brown or T.C. Maa, 1♂ (BPBM_TCN 00002956) (BPBM). brown, paler at base and darkened apically, rarely Additional material: PAPUA NEW GUINEA: yellow with pale brown apex; segment II brown Bougainville Province: Buin, 6.81361°S to dark brown, often paler basally; segments III– 155.73194°E, 29 m, 31 May 1956, J.L. Gressitt, IV dark brown. Thorax: Pronotum often yellow 1♂ (00045805) (BPBM). with pale brown or brownish anterior margin and brown posteriorly, sometimes posterior part of pronotum yellow with pale brown humeral Felisacus myersi, sp. nov. angles; mesoscutum yellow; scutellum yellow, Figures 6, 9C, 12AE, AF, 15C, 19 often pale brown to dark brown apically, some- Diagnosis: Recognized by the following times depression and punctures between mesos- combination of characters: antennal segment I cutum and scutellum pale brown; thoracic pleura ca. 1.5–1.8× as long as head width; pronotum yellow to pale brown, scent gland evaporative mostly yellow; dorsal surface of head brown to area uniformly whitish yellow to yellow, some- dark brown; inner part of clavus often whitish times with reddish or pale brown apex. Hemely- yellow to yellow; cuneus mostly colorless with tron: Mostly translucent; inner part of clavus 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 127 sometimes opaque, whitish yellow to yellow, 6D). Labium (as in Namyatova et al., 2016: figs. often with pale brown or brown apex and lateral 6D, 9C): Reaching middle of mesosternum or margins, rarely inner part of clavus pale brown slightly surpassing it; segments I and II greatly with brown to dark brown lateral margins; outer reduced, combined subequal to half of segment part of clavus whitish yellow, rarely brown api- III; segment I shorter than wide; segment II cally; corium mostly colorless, vein from apical slightly longer than wide, its dorsal surface elon- part of clavus to cuneus pale brown to brown, gate posteriorly; segment III slightly longer than often marking along inner margin on corium ventral side of head; segment IV ca. 1.5× as long pale brown to brown; embolium colorless, some- as segment III. Antenna: Segment I straight (as times whitish yellow to yellow with pale brown in Namyatova et al., 2016: fig. 8A), ca. 1.8–2.0× to brown margins; cuneus mostly colorless with as long as head width, ca. 1.2–1.3× as long as pale brown to brownish margins; membrane col- pronotum width; segment II ca. 1.9–2.3× as long orless with yellow to brown veins. Legs: Coxae as head width, ca. 1.3–1.5× as long as pronotum whitish yellow to yellow; femora whitish yellow width; segments III slightly longer than segment to yellow basally and yellow to pale brown api- II; segment IV ca. 0.3× as long as segment III. cally, sometimes also with small reddish mark- Thorax: Anterior part of pronotum slightly ings apically; tibiae yellow, often darkened basally shorter than posterior part; collar delimited; pos- and paler apically, sometimes uniformly whitish terior part slightly upraised; posterior margin of yellow to pale brown, sometimes with reddish pronotum concave; pronotum ca. 1.2–1.3× as stripe basally; tarsi uniformly pale brown to wide as long and ca. 1.4–1.6× as wide as head; brown or with segment I yellow. Abdomen: Ven- mesoscutum exposed. Hemelytron: Area along tral and lateral sides of pregenital segments whit- inner margin of corium almost flat; inner margin ish yellow, sometimes third and fourth segments of cuneus convex (as in Namyatova et al., 2016: pale brown, dorsal surface of pregenital segments fig. 13E), outer margin of cuneus ca. 3× as long reddish, pale brown or brown, genital capsule as base. Abdomen: Genital capsule rotated left at whitish yellow to pale brown. SURFACE AND right angle relative to rest of abdomen. Genitalia: VESTITURE: Corium smooth, with shallow and Genital capsule (fig. 15C) ventral wall ca. 1.7× as small punctures. Dorsum clothed with suberect long as dorsal wall, posterior margin smooth, setae slightly longer than antennal segment II semioval, without outgrowth(s), its apex inclined diameter; antennal segment I with rare suberect leftward, not curved; sides of genital capsule on setae subequal to or slightly longer than antennal modified; right paramere socket slightly acute, segment II diameter; femora with suberect setae left paramere socket rounded; distance between longer than antennal segment II diameter, abdo- paramere sockets ca. 0.4× as long as genital cap- men clothed with suberect mostly short simple sule width at base. Right paramere (fig. 12AE) setae. STRUCTURE AND MEASUREMENTS: distinctly curved in apical half; apex slightly con- Body ca. 4.5–4.6× as long as pronotum width. cave, medial part only slighter wider than basal Head: Transverse depression delimiting occipital part, bearing setae, with outer margin straight region present dorsally and laterally (as in Namy- and inner margin convex; outer angle distinct, atova et al., 2016: fig. 4E); distance between slightly swollen; inner angle indistinct, not bear- depression and pronotum distinctly shorter than ing setae; basal part of paramere ca. 0.15–0.2× as eye diameter; longitudinal sulcus on dorsal sur- long as rest of paramere. Left paramere (fig. face longer than eye diameter; distance from eye 12AF) L-shaped; apical part flattened, with tooth to pronotum slightly longer than eye diameter, on posterior side medially and without out- not swollen laterally (as in Namyatova et al., growth on dorsal surface; middle part widened, 2016: fig. 4E); vertex ca. 1.6–1.9× as wide as eye, without swelling or outgrowth; setae only on vertex upraised (as in Namyatova et al., 2016: fig. middle part near outer margin. Aedeagus (gen- 128 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 eral view as in Namyatova and Cassis, in press: semicircular sclerite and distinct sclerotized rings fig. 22I) conjunctiva weakly sclerotized, second- laterally; lateral oviducts placed almost medially, ary gonopore placed at base of vesica in repose; very close to each other, spermathecal gland placed sclerotization around secondary gonopore between lateral oviducts; dorsal labiate plate with shorter than wide; vesica with four spicules, distinct tubercles, without membranous lobe including spicules A, C, D, G, I (fig. 9C). medially. Female. Total length 3.7–4.2. COLORATION Distribution: Vanuatu (Aneityum, Tanna, (fig. 6):Head: Color pattern similar to male, but and Erromanga islands) (fig. 19). anterior side of head pale brown to brown, some- Host plants: Unknown. times with yellow markings along inferior margin Etymology: Species is names after Shepherd of antennal fossa of head; lateral side yellow to pale Myers, former curator of the BPBM insect col- brown, with brown stripe along eye margin or with lection, who kindly helped us during a visit (by brown making below eye, mandibular plate pale A.A.N.) to that museum and loaned specimens. brown or brown. Labium: Whitish yellow or yellow Discussion: Felisacus myersi is similar to F. often with segment I somewhat darker, yellow, meilingae in the external structure (fig. 6) and the brownish yellow or pale brown, segment II rarely shape of vesical spicules (cf. fig. 9B with 9C), but brownish at base, segment III rarely with reddish the latter species differs by the yellow dorsal sur- tinge. Antenna: As in male. Thorax: Additionally to face of the head, the inner part of the cuneus is coloration of pronotum described in males, collar mostly pale brown to brown and the presence of rarely brownish dorsally, posterior part of prono- additional small vesical spicule distally (fig. 9B). tum rarely almost uniformly brownish, yellow only Material examined: Holotype: VANUATU: anteriorly with dark brown markings near poste- Taeffa: Aneityum Is.: vicinity of Anelghowhat rior angles; scutellum and mesoscutum as in male, [Analgahaut], Aneityum Island, New Hebrides, but sometimes scutellum pale brown or brown; 20.22505°S 169.76858°E, 18 Jul 1971, G.F. Gross, thoracic pleura yellow to pale brown, scent gland 1♀ (00017546) (SAMA). Paratypes: VANUATU: evaporative area whitish yellow to yellow with yel- Taeffa: Aneityum Is.: Anelghowhat [Anelgao- low or pale brown apex. Hemelytron: Similar to hat], Aneityum [Aneytioum] Is., New Hebrides, male, but inner part of clavus sometimes yellow to 20.22505°S 169.76858°E, 100 m, Nov 1978, pale brown basally and brown to dark brown api- N.L.H. Krauss Coll., 1♀ (00042212) (BPBM). cally or uniformly brown to dark brown; cuneus vicinity of Anelghowhat [Analgahaut], Aneityum sometimes with yellow tinge. Legs: As in male. Island, New Hebrides, 20.22505°S 169.76858°E, Abdomen: Ventral and lateral sides of segments 18 Jul 1971, G.F. Gross, 4♀ (00017547–00017550) whitish yellow to brown; dorsal surface pale brown (SAMA); 19 Jul 1971, G.F. Gross, 1♀ (00017551), to brown. SURFACE AND VESTITURE: As in 1♂ (00017661) (SAMA); 20 Jul 1971, G.F. Gross, male. STRUCTURE AND MEASUREMENTS: 1♂ (00017662) (SAMA); 22 Jul 1971, G.F. Gross, Structure as in males; body ca. 4.3–4.6× as long as 4♀ (00017542–00017545), 6♂ (00017655– width of pronotum; vertex ca. 1.8–2.1× as wide as 00017660) (SAMA). Erromango Is.: Port Narvin eye; antennal segment I ca. 1.6–2.0× as long as head [Narevin], Erromango [Erromanga] Is., New width, ca. 1.0–1.3× as long as pronotum width; seg- Hebrides, 18.74923°S 169.20622°E, 25 Aug 1979, ment II ca. 2.0–2.1× as long as width of head, ca. N.L.H. Krauss Coll., 1♀ (00042213) (BPBM). 1.2–1.4× as long as width of pronotum; pronotum Tanna Is.: Lenakel, 0–200 m, 19.51921°S ca. 1.2–1.3× as wide as long and ca. 1.5–1.7× as 169.28217°E, 150 m, Mar 1980, N.L.H. Krauss, 1 wide as head. Genitalia (as in Namyatova et al., sex unknown (00042219) (BPBM); Jan 1981, 2016: fig. 23F, G): Dorsal labiate plate wider than N.L.H. Krauss, 1♀ (00042214) (BPBM). Lenakel, distance between apodemes of second valvula; Tanna Is., New Hebrides, 19.52733°S mostly smooth, without distinct striations, with 169.26935°E, 50 m, Nov 1978, N.L.H. Krauss 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 129
Coll., 3♂ (00042215–00042217) (BPBM). West area yellow with brown margins. Hemelytron: Coast, Tanna Is., New Hebrides, 19.54157°S Mostly translucent, colorless; inner part of cla- 169.26939°E, 12 Mar 1964, R. Straatman, 1♂ vus opaque, brown; outer part of clavus brown (00042218) (BPBM). apically; marking along inner margin of corium brown widened, not reaching R+M; embolium sometimes opaque, with brown margins; cuneus Felisacus nigrescens Carvalho sometimes opaque, with brown inner part and margins; membrane sometimes with grayish Figures 6, 9E, 13A, B, 15D, 20 tinge, with brown membrane cell. Legs: Coxa Felisacus nigrescens Carvalho, 1981: 63 (original whitish yellow; femora whitish yellow basally description). and pale brown to brown apically; tibiae pale brown to brown, often paler apically; tarsi yel- Diagnosis: Recognized by the following low to pale brown. Abdomen: Whitish ventrally combination of characters: cylindrical antennal and laterally and brown dorsally. SURFACE segment I (as in Namyatova et al., 2016: fig. 8A), AND VESTITURE: Corium smooth, with shal- transverse depression on head extending later- low and scarce punctures. Dorsum and femora ally, dorsal surface of head mostly brown; prono- clothed with suberect setae subequal to or tum brown to dark brown; marking along inner shorter than antennal segment II diameter; margin of corium brown, widened, not reaching antennal segment I with scarce suberect setae R+M; inner part of clavus brown; cuneus color- shorter than antennal segment II diameter; less or whitish with brown marking along inner abdomen clothed with suberect short simple margin; body length in male 4.4–4.7, in female setae. STRUCTURE AND MEASUREMENTS: 4.4–4.8; cuneus ca. 3× as long as base; medial Body ca. 4.4–4.7× as long as pronotum width. part of right paramere as wide as basal part, as Head: Depression, delimiting occipital region, long as basal and apical parts combined, and present dorsally and laterally (as in Namyatova with margins straight (fig. 13A); vesica with six et al., 2016: fig. 4E); distance between depres- spicules, including spicule A (fig. 9E). sion and pronotum distinctly shorter than eye Redescription: Male. Total length 4.4–4.7. diameter; longitudinal sulcus on dorsal surface COLORATION (fig. 6): Head: Pale brown to longer than eye diameter; distance from eye to brown, longitudinal sulcus dark brown; anterior pronotum slightly longer than eye diameter, not side yellow to pale brown, sometimes with red- swollen laterally (as in Namyatova et al., 2016: dish tinge; clypeus yellow to pale brown basally fig. 4E); vertex ca. 1.5–1.7× as wide as eye, and brown apically, sometimes with reddish upraised (as in Namyatova et al., 2016: fig. 6D). marking, tubercles around antennal fossa pale Labium (as in Namyatova et al., 2016: figs. 6D, brown; lateral sides and buccula pale brown, 9C): Almost reaching posterior margin of mandibular and maxillary plates whitish yellow mesosternum; segments I and II combined to yellow. Eye brown with reddish tinge or dark shorter than half of segment III; strongly brown. Labium: Uniformly yellow. Antenna: reduced; segment I shorter than wide; segment Segments I dark brown; segment II brown to II slightly longer than wide, its dorsal surface dark brown, sometimes pale brown apically, elongate posteriorly; segment III slightly longer segments III–IV brown, segment III darkened than ventral side of head; segment IV ca. 1.5× basally. Thorax: Pronotum brown to dark as long as segment III. Antenna: Segment I brown, somewhat paler medially; scutellum and cylindrical (as in Namyatova et al., 2016: fig. mesoscutum yellow to pale brown, sometimes 8A), ca. 1.3–1.6× as long as head width, ca. mesoscutum yellow and scutellum pale brown; 0.9–1.0× as long as pronotum width; segment II thoracic pleura brown; scent gland evaporative ca. 1.6–1.9× as long as head width, ca. 1.0–1.2× 130 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 as long as pronotum width; segment III slightly TURE AND MEASUREMENTS: Structure as in longer than segment II. Thorax: Anterior part of male; body ca. 4.3–4.9× as long as pronotum pronotum shorter than posterior part; collar width; vertex ca. 1.9–2.1× as wide as eye; anten- delimited; posterior part slightly upraised; pos- nal segment I ca. 1.4–1.5× as long as head width, terior margin of pronotum concave; pronotum ca. 0.8–0.9× as long as pronotum width; segment ca. 1.3–1.4× as wide as long and ca. 1.4–1.7× as II ca. 1.7–1.9× as long as head width, ca. 1.0– wide as head; mesoscutum exposed. Hemely- 1.1× as long as pronotum width; segment IV ca. tron: Area along inner margin of corium flat; 0.3× as long as segment III; pronotum ca. 1.3× as inner margin of corium concave (as in Namya- wide as long and ca. 1.7× as wide as head. Geni- tova et al., 2016: fig. 13E), outer margin ca. 3× talia (as in Namyatova et al., 2016: fig. 23F, G): as long as base. Abdomen: Genital capsule Dorsal labiate plate wider than distance between rotated left at right angle relative to rest of apodemes of second valvula; mostly smooth, abdomen. Genitalia: Genital capsule (fig. 15D) without distinct striations, with semicircular ca. 1.5× as long as wide; ventral wall ca. 1.5× as sclerite and distinct sclerotized rings laterally; long as dorsal wall, its posterior margin smooth, lateral oviducts placed almost medially, very semioval, without outgrowth(s), not curved; its close to each other, spermathecal gland placed apex inclined rightward; sides of genital capsule between lateral oviducts; dorsal labiate plate with not modified; margins of paramere sockets distinct tubercles, without membranous lobe more or less rounded; distance between param- medially. ere sockets ca. 0.7× as long as genital capsule Distribution: Papua New Guinea (fig. 20). width at base. Right paramere (fig. 13A) dis- Host plants: Pteris gardneri (Pteridaceae— tinctly curved in apical half; apex distinctly large number of specimens; Ptelea sp. (Ruta- concave posteriorly; medial part only slighter ceae)—single specimen; beating ferns wider than basal part, bearing setae; outer and (Carvalho, 1981). inner margins of middle part almost straight; Discussion: Segment IV is lost in male speci- outer angle distinct; inner angle rounded, with- mens that we examined. Carvalho (1981) out setae; basal part of paramere ca. 0.15–0.2× described F. nigrescens on the basis of male and as long as rest of paramere. Left paramere (fig. female specimens. We examined the paratypes of 13B) L-shaped; apical part not flattened, with both sexes. Based on the coloration and the vesi- tooth on posterior side medially (as in fig. 11G) cal spicules, Felisacus nigrescens is not similar to and without outgrowth on dorsal surface; mid- any other species of the genus. dle part widened, without swelling or out- Material examined: Paratypes: PAPUA NEW growth; setae only on middle part near outer GUINEA: Eastern Highlands: NE Okapa, margin. Aedeagus (general view as in Namya- 6.56475°S 145.60089°E, 1900 m, 24 Jun 1967–25 tova and Cassis, in press: fig. 22I) conjunctiva Jun 1967, G.A. Samuelson, 1♀ (00043247) (BPBM), weakly sclerotized; secondary gonopore placed 2♀ (00018981, 00018982) (TAMU). Morobe Prov- at base of vesica in repose; sclerotization of duc- ince: Wau, 7.3333°S 146.71667°E, 1750 m, 06 Sep tus seminis around secondary gonopore shorter 1965, J. and M. Sedlacek, 1♂ (00043244) (BPBM). than wide; vesica with six spicules; including Additional material: PAPUA NEW GUINEA: spicule A (fig. 9E). Morobe Province: Edie Creek, NE Wau, 7.31666°S Female. Total length 4.4–4.8. COLORATION 146.68333°E, 1100 m, 08 Sep 1971, W.C. Gagne, (fig. 6): Similar to male, but mesoscutum some- 1♂ (00043245) (BPBM). Mt. Missim, 7.1167°S times reddish, legs rarely with reddish tinge; 146.9167°E, 1600 m, 23 Mar 1978, W.C. Gagne, 1♀ abdomen whitish yellow laterally and ventrally, (00045819) Pteris gardneri (Pteridaceae), 1♂ with reddish segment X, brown dorsally. SUR- (00043246), 2♀ (00043248, 00043249), 2 sex FACE AND VESTITURE: As in male. STRUC- unknown (00043250, 00043251) (BPBM). Namie 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 131
Creek, 7.31666°N 146.7°E, 25 May 1982, P. Groot- Distribution: New Guinea (fig. 21). aert, Ptelea sp. (Rutaceae), 1♂ (00018973) (ISNB). Host plants: Unknown. Discussion: The type of Felisacus nigricornis Felisacus nigricornis Poppius is preserved in the Hungarian Museum of Natu- Figure 21 ral Sciences, but is mutilated (Carvalho, 1980); this was also confirmed by the curators of this Felisacus nigricornis Poppius, 1912: 2 (original collection (András Orosz and Dávid Rédei, per- description). sonal commun.). We did not find any additional Diagnosis: Recognized by the following specimens which could be assigned to this spe- characters: head, pronotum, scutellum, base of cies. Poppius (1911) compared F. nigricornis with clavus red; main part of clavus, claval commis- F. elegantulus. Based on Poppius’ description, the sure apically, inner margin of corium and anten- affinities of this species are not clear. Carvalho nal segments I–II black; outer and inner parts of (1981) also redescribed F. nigricornis based on cuneus brown; body length 4.0. additional material. Description (from Poppius (1912), trans- There are six Felisacus species known from lated from German): New Guinea: F. dauloi, F. javanus, F. nigres- cens. F. longiceps, F. magnificus, and F. minu- Head, pronotum, scutellum, as well as base of clavus, tus. They all differ from F. nigricornis in red; apex of clypeus, collar, rest of clavus, claval com- coloration. Felisacus dauloi can be separated missure apically, inner margin of corium, narrow from F. nigricornis in the head, pronotum, outer margin of embolium, as well as antennal seg- scutellum, and inner part of the clavus mostly ments I–II (segments III–IV lost) black; narrow inner yellow, the outer part of the clavus colorless, margin of embolium, corium apically, outer and antennal segment I–II reddish brown, the inner parts of cuneus, as well as membrane cell, cuneus mostly colorless, translucent with its brown; rest of hemelytron transparent, colorless; ven- margins and often apex yellow, red or mostly tral side of body red, posterior part of body with yel- yellow. Felisacus javanus differs from F. nigri- low marking medially; antennal segment I basally cornis by the head yellow to brown, antennal red; labium and legs yellow; femora basally whitish segment I pale brown, sometimes with a red- yellow, somewhat reddish apically. Head from ante- dish tinge, segment II reddish brown, the rior view slightly longer than wide, tapering; occipital pronotum mostly yellow, the scutellum whit- region delimited with depression; from lateral view ish yellow to pale brown, sometimes with a slightly longer than basal height; frons slightly less reddish tinge laterally, the inner part of the than twice as long as eye diameter; eye rather large, clavus mostly whitish yellow to yellow, the distinctly protruding, very finely granulate; labium outer part of the clavus colorless; the cuneus reaching mesofemora, with segment I not reaching with its inner part red and the outer part posterior margin of head; segment II slightly longer, whitish yellow. Felisacus nigrescens differs slightly narrower than segment I. Antennae long, from F. nigricornis by the head mostly pale clothed with suberect setae; segment I almost longer brown, brown dorsally, the pronotum brown than pronotum, segment II slightly longer, slightly to dark brown, the scutellum yellow to pale narrower than segment I. Pronotum distinctly longer brown, the inner part of the clavus brown, the than wide, posterior margin almost twice as long as outer part of the clavus colorless, brown api- anterior margin; calli delimited with depression; calli cally, the cuneus colorless with the inner part large, merged with each other, collar narrow; prono- and margins brown. Felisacus minutus is sep- tal disc distinctly convex and declivous. Hemelytron arated from F. nigricornis in the head and the in female longer than posterior part of body. Legs anterior part of the pronotum mostly pale with pale setae. Length 4, width 1 mm. brown with a reddish tinge, the posterior part 132 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 of the pronotum brown with a wide yellow reddish brown. Thorax: Pronotum whitish yellow stripe medially, the cuneus red with the outer to yellow, anterior margin and punctures between part yellow and the margins brown. Felisacus anterior and posterior parts pale brown; scutel- magnificus and F. longiceps differ from F. lum and mesoscutum whitish yellow to yellow, nigricornis by the head and the anterior part depression between them pale brown; thoracic of the pronotum yellow to pale brown, the pleura yellow, scent gland evaporative area whit- posterior part of the pronotum dark brown to ish yellow, paler than thoracic pleura, or whitish black, the C-shaped marking on the corium basally and yellow or reddish apically. Hemely- reaching R+M anteriorly and posteriorly, and tron: Mostly translucent and colorless; inner part the cuneus mostly colorless and red apically. of clavus whitish yellow, with pale brown or No other Felisacus species have the head, pro- brown margins; corium with marking along api- notum, and scutellum red. cal part of clavus pale brown, marking along inner margin of corium yellow, sometimes red Felisacus ochraceus Usinger posteriorly; embolium with yellow or red apex Figures 6, 9F, 13C, D, 15E, 19 and yellow or pale brown margins; cuneus with yellow or reddish inner part, margins yellow to Felisacus ochraceus Usinger, 1946: 69 (original pale brown; sometimes cuneus uniformly red- description). dish yellow; membrane cell yellow. Legs: Mostly Diagnosis: Recognized by the following whitish yellow, femora apically and tibiae basally combination of characters: cylindrical antennal yellow, sometimes femora apically and entire segment I (as in Namyatova et al., 2016: fig. 8A), tarsi reddish yellow, tarsi yellow to pale brown. transverse depression on head extending later- Abdomen: Yellow with red markings dorsally and ally, vertex upraised (as in Namyatova et al., laterally or uniformly yellow. SURFACE AND 2016: fig. 6D); main coloration of head and pro- VESTITURE: Corium smooth, shallow and notum yellow to pale brown, humeral angles not scarce punctures. Dorsum, antennal segment I darkened; inner part of clavus whitish yellow, and femora clothed with setae shorter than with pale brown or brown margins; cuneus with antennal segment II diameter; abdomen clothed whitish and translucent outer part and yellow or with short erect setae. STRUCTURE AND MEA- reddish inner part; labium reaching posterior SUREMENTS: Body ca. 4.3–4.7× as long as pro- margin of mesosternum; cuneus 3× as long as notum width. Head: Depression delimiting wide; medial part of right paramere as wide as occipital region distinct dorsally and laterally (as basal part, subequal to basal and apical parts in Namyatova et al., 2016: fig. 4E), distance combined, its outer margin concave and inner between depression and pronotum distinctly margin convex (fig. 13C); vesica with five spic- shorter than eye diameter; longitudinal sulcus on ules, including spicules A, B, C, and H, spicule B dorsal surface of head as long as eye diameter; with long distal arm (fig. 9F). distance from eye to pronotum longer than eye Redescription: Male. Total length 3.6–4.0. diameter, not swollen laterally (as in Namyatova COLORATION (fig. 6): Head: Mostly yellow, et al., 2016: fig. 4E); vertex ca. 1.4–1.7× as wide sometimes with marking near antennal fossa and as eye, upraised (as in Namyatova et al., 2016: fig. stripe behind eye reddish, dorsal surface often 6D). Labium: Reaching posterior margin of with reddish tinge, longitudinal sulcus some- mesosternum or slightly surpassing it; segments times pale brown. Eye dark brown with reddish I and II strongly reduced, combined shorter than tinge. Labium: Uniformly whitish yellow to yel- half of segment III; segments I as long as wide low. Antenna: Segment I yellow basally and pale (as in Namyatova et al., 2016: figs. 6D, 9C); seg- brown apically, sometimes reddish yellow; seg- ment II slightly longer than wide, its dorsal sur- ments II–III pale brown to brown, sometimes face elongate posteriorly; segment III longer than 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 133 ventral side of head; segment IV ca. 1.5× as long Female. Total length 3.8–3.9. COLORATION as segment III. Antenna: Segment I cylindrical (fig. 6): Head: Similar to male, but dorsal surface (as in Namyatova et al., 2016: fig. 8A), ca. 1.5– slightly darker than other sides. Labium: As in 1.6× as long as head width, ca. 1.0–1.1× as long male. Antenna: Similar to male, segment II uni- as pronotum width; segment II twice as long as formly reddish brown. Thorax and legs: As in head width, ca. 1.4–1.5× as long as pronotum male. Hemelytron: Similar to male, but cuneus width; segment III slightly longer than segment with outer part colorless and with inner part II. Thorax: Anterior and posterior parts subequal reddish or pale brown, rarely mostly yellow with in length, collar delimited; posterior part slightly pale brown inner part. Abdomen: Pale brown upraised; posterior margin concave, pronotum with reddish dorsal surface. SURFACE AND ca. 1.2–1.3× as wide as long and ca. 1.4–1.5× as VESTITURE: As in male. STRUCTURE AND wide as head; mesoscutum exposed; Hemelytron: MEASUREMENTS: Structure as in male; body Area along inner margin of corium flat; inner ca. 4.2–4.4× as long as pronotum width; ca. ver- margin of cuneus convex (as in Namyatova et al., tex 1.3–1.6× as wide as eye; antennal segment I 2016: fig. 13E), outer margin ca. 3× as long as ca. 1.2–1.5× as long as head width, ca. 0.9–1.0× base. Abdomen: Genital capsule rotated left at as long as pronotum width; segment II ca. 1.9– right angle relative to rest of abdomen. Genitalia: 2.0× as long as head width, ca. 1.3–1.4× as long Genital capsule (fig. 15E) ca. 1.5× as long as as pronotum width; pronotum ca. 1.1–1.3× as wide; ventral wall ca. 1.5× as long as dorsal wall, wide as long and ca. 1.4–1.6× as wide as head. with posterior margin of ventral wall semioval, Genitalia (as in Namyatova et al., 2016: fig. 23F, smooth, without outgrowth(s), its apex inclined G): Dorsal labiate plate wider than distance leftward, not curved dorsally; sides of genital between apodemes of second valvula; mostly capsule not modified; right paramere socket smooth, without distinct striations, with semi- slightly angulate and left socket rounded; dis- circular sclerite and distinct sclerotized rings tance between paramere sockets subequal to half laterally; lateral oviducts placed almost medially, of genital capsule width at base; right angle of left very close to each other, spermathecal gland paramere socket not projecting. Right paramere placed between lateral oviducts; dorsal labiate (fig. 13C) distinctly curved in apical half; apex plate with distinct tubercles, without membra- slightly concave; medial part as wide as basal nous lobe medially. part, bearing setae, with outer margin concave Distribution: Guam (fig. 19). and inner margin convex; outer angle distinct; Host plants: Ferns (Usinger, 1946). inner angle indistinct, without setae; basal part Discussion: Antennal segment IV is lost in very short, basal part of paramere ca. 0.15–0.2× males and segment III–IV are lost in females. as long as rest of paramere. Left paramere (fig. Usinger (1946) described F. ochraceus from a 13D) L-shaped, apical part flattened, with tooth- number of specimens. We examined the paratypes like outgrowth on posterior side medially (as in of both sexes. Based on the structure of the para- fig. 11G) and without outgrowth on dorsal sur- meres, the specimens described by Carvalho face; middle part widened, without swelling or (1981) as F. ochraceus belongs to another species, outgrowth; setae only on middle part near outer whose identity we were not able to establish; how- margin. Aedeagus (general view as in Namyatova ever, the shape of parameres in figures 115 and et al., 2016: fig. 22I) conjunctiva weakly sclero- 116 of Carvalho (1981) is similar to those of F. tized, sclerotized part of ductus seminis very nigrescens and F. solomonicus (figs. 13A, R), but short, secondary gonopore placed at base of coloration differs from both of these species. vesica in repose; vesica with five spicules, includ- Felisacus ochraceus is almost identical with F. ing spicules A, B, C, and H, spicule B with long australicus in structure, including the parameres apical arm (fig. 9F). (cf. fig. 11E, F with fig. 13C, D) and vesical spic- 134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 ules (cf. fig. 8C with 9F), but the latter species dif- rounded outgrowths (fig. 13F); sclerotization fers in coloration, with the head, antennal segment around ductus seminis distinctly elongate, sec- I, and the pronotum mostly yellow to pale brown, ondary gonopore placed at base of vesica in the inner part of clavus pale brown to brown, and repose; vesica with a number of small toothlike the scent gland evaporative area pale brown. spinules and two large sclerites (fig. 9G). Felisacus ochraceus is also similar in coloration Description: Male. Total length 3.2. COLOR- to F. amboinae, F. filicicola, F. dauloi, F. insularis, ATION (fig. 6): Head: Mostly yellow with reddish and F. linae (figs. 4, 5). All these species can be tinge, mandibular and maxillary plates whitish separated from F. ochraceus by the medial part of yellow. Eye dark brown with reddish tinge. right paramere wider than the basal part and dis- Labium: Uniformly yellow. Antenna: Segment I tinctly shorter than the basal and apical parts yellow with reddish tinge, whitish yellow basally combined, with the outer margin straight or and reddish apically; segment II uniformly red- slightly concave (figs. 11C, X, AD, 12C, K), as well dish. Thorax: Pronotum yellow with brown mark- as in the number and shape of the vesical spicules ing near humeral angle; scutellum and (fig. 8B, G, I, L, P ). Felisacus dauloi also differs mesoscutum yellow with reddish tinge; thoracic from F. ochraceus by the cuneus colorless or yel- pleura yellow; scent gland evaporative area whit- low basallyand reddish apically. Felisacus linae dif- ish yellow, with reddish marking apically. Hemely- fers from F. ochraceus by the cuneus mostly tron: Mainly whitish yellow, translucent; clavus translucent with the inner part pale brown. yellow, outer part paler than inner part; marking Material examined: Paratypes: GUAM: along inner margin of corium yellow to pale Piti, 13.46°N 144.69166°E, 5 m, 24 Aug 1936, brown, narrow, not reaching R+M; embolium O.H. Swezey, 1♂ 00399747), 3♀ (00399751, whitish yellow, yellow apically; cuneus whitish 00399749, 00399750) (CAS). Additional material: with yellow tinge; membrane translucent with GUAM: Mariana Is: Guam, Mt. Lamlam, Oct brown tinge, membrane cell pale brown. Legs: 1957, N.L.H. Krauss, 2♂ (00043157, 00043159) Mostly whitish yellow; apical parts of femora yel- (BPBM). Mt. Alifan, 13.38055°N 144.67138°E, low with reddish tinge, tibiae yellow with reddish Oct 1957, N.L.H. Krauss, 1♀ (00043161) tinge basally and apically; tarsi brown. SURFACE (BPBM). Yigo, 13.53194°N 144.88027°E, 126 m, AND VESTITURE: Corium with distinct puncta- Oct 1957, N.L.H. Krauss, 1♂ (00043158), 2♀ tion. Dorsum with setae subequal to or slightly (00043160, 00043162) (BPBM). longer than antennal segment II diameter; anten- nal segment I with suberect setae shorter than antennal segment II diameter; femora with suber- Felisacus ovalau, sp. nov. ect setae longer than antennal segment II diame- Figures 6, 9G, 13E, F, 15F, 19 ter; abdomen clothed with short erect setae. Diagnosis: Recognized by the following STRUCTURE AND MEASUREMENTS: Body ca. combination of characters: transverse depres- 4.6× as long as pronotum width. Head: Depres- sion on head present only dorsally; head and sion delimiting occipital region present only dor- pronotum mostly yellow; antennal segment I sally; distance between depression and pronotum yellow with reddish tinge; posterior part of pro- distinctly shorter than eye diameter; longitudinal notum yellow with pair of brown marking near sulcus on dorsal surface of head shorter than eye humeral angles; labium reaching abdominal diameter; distance from eye to pronotum longer segment V; inner margin of corium convex (as than eye diameter, not swollen laterally; vertex ca. in Namyatova et al., 2016: fig. 13E); apex of ven- 2.0× as wide as eye, upraised. Labium (as in tral wall of genital capsule slightly curved dor- Namyatova et al., 2016: figs. 6D, 9C): Reaching sally (fig. 15F); right paramere sickle shaped abdominal sternum V; segments I and II com- (fig. 13E); left paramere wide, with three bined longer than half of segment III; segment I 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 135 shorter than width; segment II twice as long as Female. Unknown. wide, its dorsal surface elongate posteriorly; seg- Distribution: Tahiti (fig. 19). ment III longer than ventral side of head; segment Host plants: Unknown. IV ca. 0.3× as long as segment III. Antenna: Seg- Etymology: The species is named after Ova- ment I distinctly longer than head width, slightly lau Is. (Fiji), where it was collected. widened, ca. 1.4× as long as head width, 0.9× as Discussion: Felisacus ovalau is similar to F. long as pronotum width; segment II ca. 1.8× as bradi, F. usingeri, and F. webbi in coloration and long as head width, 1.3× as long as pronotum structure (figs. 4, 7). Felisacus bradi can be sepa- width; antennal segments III–IV lost. Thorax: rated from F. ovalau in the antennal segment I Anterior part of pronotum somewhat shorter than mostly brown, the pronotum without brown posterior part, collar delimited; posterior part of markings, the left paramere with a single out- pronotum slightly upraised; posterior margin con- growth posteriorly (fig. 11O), the sclerite around cave, pronotum ca. 1.1× as wide as long and ca. the ductus seminis distinctly longer than wide 1.5× as wide as head; mesoscutum exposed. Hem- (fig. 10B). Felisacus webbi differs by from F. ova- elytron: Area along inner margin of corium flat; lau by the ventral wall of the genital capsule not inner margin of corium convex (as in Namyatova curved apically, and with an outgrowth dorsally et al., 2016: fig. 13E), its outer margin ca. 3× as and to the right-hand side (fig. 15N), the left long as base. Abdomen: Genital capsule rotated paramere sickle shaped, without outgrowths (fig. left at right angle relative to rest of abdomen. Gen- 13AA), and the vesica has a long curved spicule, italia: Genital capsule (fig. 15F) ca. 1.5× as long as and without toothlike spinules (fig. 10N). Feli- wide; ventral wall ca. 1.5× as long as dorsal wall, sacus usingeri differs from F. ovalau by antennal with posterior margin of ventral wall acute api- segment I yellow, brown apically, the pronotum cally, smooth, without outgrowth, its apex placed without brown markings, and the labium reaches almost medially, curved dorsally; right side of the middle of the mesosternum. genital capsule not curved, left side not widened; Felisacus vitilevu also inhabits Fiji and is right paramere socket acute, left one angulate; dis- closely related to F. ovalau. However, F. vitilevu tance between paramere sockets ca. 0.3× as long differs in the pronotum mostly dark brown with as genital capsule width at base. Right paramere a yellow stripe medially (fig. 7), the left paramere (fig. 13E) distinctly curved in apical half; apex without a rounded outgrowth, only with a single concave posteriorly; medial part narrower than outgrowth posteriorly (fig. 13W), and sclerotiza- basal part with setae near inner margin, its outer tion of the ductus seminis around the secondary margin convex and inner margin concave; param- gonopore longer than wide (fig. 9N). ere slightly swollen dorsally; outer and inner Material examined: Holotype: FIJI: Cen- angles rounded, not bearing setae; basal part of tral Division: Levuka, Ovalau Is., 17.68039°S paramere ca. 0.4× as long as rest of paramere. Left 178.82994°E, 100 m, Dec 1978, N.L.H. Krauss, paramere (fig. 13F) irregularly shaped; apical part 1♂ (00042352) (BPBM). curved, flattened, without outgrowth posteriorly; middle part with three rounded swellings, setae Felisacus philippinensis Hsiao placed on middle part near outer margin. Aedeagus conjunctiva sclerotized basally and with Figures 6, 10K, 13G, H, 15G, 16G. H, 20 sclerotized teeth apically; secondary gonopore place at base of vesica in repose, sclerotization of Felisacus philippinensis Hsiao, 1944: 386 (original ductus seminis around secondary gonopore as description). long as wide, its dorsal wall distinctly longer than ventral part; vesica with teeth apically and two Diagnosis: Recognized by the following com- spicules (fig. 9G). bination of characters: dorsal side of head and 136 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 pronotum mostly yellow; marking along inner yellow tinge, outer margin yellow and inner mar- margin of corium yellow to dark brown, not con- gin whitish; membrane colorless, sometimes pale tinuing toward R+M (fig. 6); transverse depres- brown to brown medially; membrane cell pale sion on head not continuing laterally; antennal brown to dark brown. Legs: Often uniformly yel- segment I widened (as in Namyatova and Cassis, low, sometimes femora apically and tibiae basally in press: fig. 8B), red; labium slightly surpassing red, sometimes middle tibia and fore- and mid- posterior margin of prosternum; inner margin of dle tarsal segments II–III pale brown to brown. cuneus straight (as in Namyatova et al., 2016: fig. Abdomen: Dorsal and lateral sides mostly yellow, 13F); genital capsule widened from left-hand side often with brown markings and stripe, genital dorsally, its ventral wall not curved apically (fig. capsule sometimes with reddish tinge, dorsal 15G); right paramere sickle shaped, with outer surface red or brown. SURFACE AND VESTI- angle indistinct and inner angle distinct (fig. 13G); TURE: Corium with shallow punctation. Dor- apical half of ductus seminis sclerotized, hooked sum and femora with setae subequal to or slightly apically (fig. 15G). longer than antennal segment II diameter; anten- Redescription: Male. Total length 3.9–4.1. nal segment I clothed with adpressed setae COLORATION (fig. 6): Head: Mostly yellow, shorter than antennal segment II diameter; often with pale brown or red markings; tubercles abdomen clothed with erect setae of different around antennal fossa whitish yellow; clypeus length. STRUCTURE AND MEASUREMENTS: reddish to brown with yellow base; mandibular Body ca. 4.4–4.6× as long as pronotum width. and maxillary plates yellow, buccula often yellow, Head: Depression delimiting occipital region brown to dark brown anteriorly, rarely uniformly present only dorsally; distance between depres- yellow or brown to dark brown. Eye dark brown sion and pronotum as long as eye diameter, not with reddish tinge. Labium: Segments I–II pale swollen laterally; longitudinal sulcus on dorsal brown to brown, segments III–IV pale brown to surface of head shorter than eye diameter; dis- brown, segment III often paler than segment IV. tance from eye to pronotum longer than eye Antenna: Segment I reddish, sometimes reddish diameter, not swollen laterally; vertex ca. 1.4– brown; segment II reddish brown, segments III– 1.6× as wide as eye; upraised. Labium: Slightly IV brown to dark brown. Thorax: Pronotum surpassing prosternum; segments I and II longer mostly yellow, anterior margin often pale brown than wide, II combined as long as segment III; to brown; collar and calli often with reddish dorsal surface of segment II not elongate poste- tinge; posterior margin often pale brown with riorly (as in Namyatova et al., 2016: fig. 6E); seg- pair of brown to brown markings near humeral ment III distinctly shorter than ventral side of angles, sometimes uniformly yellow to brown; head; segment IV ca. 1.5× as long as segment III. scutellum and mesoscutum yellow to pale brown, Antenna: Segment I widened basally and medi- sometimes with reddish tinge; thoracic pleura ally (as in Namyatova et al., 2016: fig. 8B), ca. usually pale brown, sometimes mostly brown to 1.3–1.6× as long as head width, ca. 0.9× as long dark brown; scent gland evaporative area uni- as pronotum width; segment II ca. 1.8–2.2× as formly whitish yellow to yellow. Hemelytron: long as head width, ca. 1.2–1.3× as long as pro- Mostly translucent; clavus sometimes opaque, its notum width; antennal segment III slightly lon- inner part pale brown to dark brown, rarely yel- ger than segment II, segment IV only slightly low; outer part mostly pale brown to dark brown, shorter than segment III. Thorax: Anterior part rarely colorless or yellow, often paler than inner of pronotum distinctly shorter than posterior part; corium with yellow tinge; marking along part; collar distinct; posterior part slightly inner margin of corium yellow to dark brown; upraised; posterior margin of pronotum straight embolium colorless, with yellow tinge and yellow or concave; pronotum ca. 1.1–1.2× as wide as to pale brown margins; cuneus colorless, with long and ca. 1.4–1.9× as wide as head; mesoscu- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 137 tum exposed. Hemelytron: Area along inner mar- MENTS: Structure as in male; body ca. 4.5–4.7× gin of corium swollen; inner margin of cuneus as long as pronotum width; vertex ca. 1.4–1.6× as straight (as in Namyatova et al., 2016: fig. 13F), wide as eye; antennal segment I ca. 1.3–1.5× as outer margin of cuneus twice as long as width at long as head width, ca. 0.8× as long as pronotum base. Abdomen: Genital capsule rotated left at width; segment II ca. 1.9–2.1× as long as head right angle relative to the rest of abdomen. Geni- width, ca. 1.1–1.2× as long as pronotum width; talia: Genital capsule (fig. 15G) ca. 1.5× as long pronotum ca. 1.1–1.2× as wide as long and ca. as wide; ventral wall ca. 1.5× as long as dorsal 1.8–1.9× as head wide. Genitalia (figs. 16G, H): wall, with posterior margin smooth, without Dorsal labiate plate very small and transparent, as outgrowth(s), not curved; right side of genital wide as distance with apodemes of second valvu- capsule not curved, left side widened; paramere lae, without striations or membranous ridge sockets angulate; distance between paramere medially; semicircular sclerite and sclerotized sockets ca. ca. 0.25–0.3 as long as width of geni- rings absent; lateral oviducts placed in posterior tal capsule at base. Right paramere (fig. 13G) part, close to each other, far from lateral margins distinctly curved apically; apex concave posteri- of dorsal labiate plate; spermathecal gland attached orly, medial part narrower than basal part, with- near anterior margin; dorsal labiate plate without out setae, with outer margin convex and inner distinct tubercles, with triangular membranous margin widened; outer angle indistinct, rounded, lobe posteriorly. inner angle distinct, widened, bearing setae; Distribution: Philippines (Luzon, Mind- basal part of paramere subequal to half of rest of anao, Negros) (fig. 20). Only females are known paramere. Left paramere (fig. 13H) irregularly from Negros. shaped; apical part of paramere flattened and Host plants: Unknown. widened, with four outgrowths medially of dif- Discussion: Hsiao (1944) described F. philip- ferent shape and length; setae placed on middle pinensis, and the types are preserved in the part at inner margin. Aedeagus conjunctiva USNM and were examined but not dissected. We membranous; vesica without spicules; apical half have specimens which fit the description of of ductus seminis sclerotized, hooked apically, Hsiao and we treated them as conspecific with F. secondary gonopore placed near phallotheca philippinensis. mouth in repose (fig. 10K). Felisacus philippinensis is similar to F. luzonus Female. Total length 4.3–4.9. COLORATION externally (figs. 5, 6), but the latter species differs (fig. 6): Head: Similar to male, but dorsal surface by the posterior part of the marking along the sometimes mostly reddish brown. Labium: Simi- corium extends, to R+M, the labium reaches the lar to male, segment III sometimes pale brown to middle of the mesosternum, the genital capsule brown, of the same color as segments I–II. is not broad on the right-hand side (fig. 14X) and Antenna: As in male. Thorax: Similar to male, but the ductus seminis is straight apically (fig. 10I). pronotum yellow to pale brown with brown to Material examined: PHILIPPINES: Cama- dark brown lateral sides, scutellum yellow to rines Sur: Iriga Mt, 13.45416°N 123.45055°E, 26 brown, sometimes with reddish tinge. Hemelytron: Apr 1962, H.M. Torrevillas, 2♂ (00043115, Similar to male. Legs: Similar to male, but some- 00043116) (BPBM). Iriga Mt, 13.45416°N times coxae and femora basally whitish yellow and 123.45055°E, 550 m, 24 Apr 1962, H.M. Torrevil- tibiae yellow basally and whitish yellow apically. las, 1♂ (00043114), 2♀ (00043119, 00043120) Abdomen: Ventral and lateral sides mostly yellow, (BPBM). Isarog Mt, 20 km E of Naga, 13.65805°N often with brown stripes and markings, segment 123.37305°E, 550 m, 06 Apr 1963, H.M. Torrevil- IX often brown or reddish brown, dorsal surface las, 1♀ (00043112) (BPBM). Isarog Mt, Pili, brown or reddish. SURFACE AND VESTITURE: 13.65805°N 123.37305°E, 850 m, 12 Apr 1965, As in male. STRUCTURE AND MEASURE- L.M. Torrevillas, 8♂ (00043097–00043104), 2♀ 138 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
(00043105, 00043107) (BPBM). Isarog Mt, Pili, Description: Male. Total length 3.5–3.6. 13.65805°N 123.37305°E, 600 m, 20 Sep 1964, COLORATION (fig. 6):Head: Mostly yellow. M.D. Delfinado, 2♀ (00043109, 00043110) Eye dark brown with reddish tinge. Labium: (BPBM). Isarog Mt, Pili, 13.65805°N 123.37305°E, Uniformly yellow. Antenna: Segment I yellow, 500 m, 04 Apr 1963, H.M. Torrevillas, 1♂ sometime with reddish tinge, segment II red, (00043113) (BPBM). Isarog Mt, Pili, 13.65805°N segments III–IV reddish brown. Thorax: Often 123.37305°E, 100 m, 20 Apr 1965, H.M. Torrevil- uniformly yellow, pronotum and scutellum las, 1♀ (00043108) (BPBM). Mt. Isarog, Pili, sometimes with reddish tinge dorsally. thoracic 13.70805°N 123.75083°E, 800 m, 27 Apr 1965, pleura and scent gland evaporative area whitish H.M. Torrevillas, 1♀ (00043111) (BPBM). yellow to yellow. Hemelytron: Mainly whitish Davao del Sur: Galog Riv., Mt. Apo, Mindanao, yellow, translucent; clavus yellow, inner part 1829 m, 07 Sep 1700, C.S. Clagg, 1♂ (00045807) opaque; marking along inner margin of corium (BPBM). Mindanao: Matibog Creek, Bayugan, yellow to pale brown; embolium whitish yellow, Esperanza, Agusan, 8.91716°N 125.78854°E, 300 opaque, yellow to pale brown apically; cuneus m, 07 Nov 1959, C.M. Yoshimoto, 1♂ (00043128) whitish with yellow tinge, sometimes opaque; (BPBM). Negros Oriental: Balinsasayao Lake, membrane with brown tinge, membrane cell 9.35305°N 123.17916°E, 01 Oct 1959–07 Oct yellow to brown. Legs: Whitish yellow to pale 1959, L.W. Quate, 2♀ (00043123, 00043125) brown, tarsi sometimes pale brown. SURFACE (BPBM); 01 Oct 1959–07 Oct 1959, C.M. Yoshi- AND VESTITURE: Corium smooth without moto, 3♀ (00043124, 00043126, 00043127), 1 punctation. Dorsum, femora, and abdomen juvenile (00043121) (BPBM). Surigao: Lake with setae longer than antennal segment II Mainit, 9.4333°N 125.5333°E, 24 Nov 1959–01 diameter; antennal segment I with suberect Dec 1959, L.W. Quate, 3♂ (00043122, 00043117, setae shorter than antennal segment II diame- 00043118) (BPBM). ter. STRUCTURE AND MEASUREMENTS: Body ca. 3.5–3.7× as long as pronotum width. Head: Depression delimiting occipital region Felisacus ponaponensis (Carvalho) present dorsally and laterally; distance between Figures 6, 10L, 13I, J, 15H, 20 depression and pronotum distinctly shorter Felisacoris ponaponensis Carvalho, 1956: 24 than eye diameter; longitudinal sulcus on dorsal (original description), comb. nov. surface of head shorter than eye diameter; dis- tance from eye to pronotum shorter than eye Diagnosis: Recognized by the following com- diameter, not swollen laterally; vertex ca. 1.5– bination of characters: collar not delimited; ante- 1.8× as wide as eye, upraised; head slightly rior margin of pronotum distinctly raised and swollen dorsally. Labium: Reaching or almost covering head, almost reaching eyes; mostly yel- reaching posterior margin on prosternum; seg- low coloration, marking along inner margin of ments I and II combined subequal to segment corium yellow to pale brown, narrow, not extend- III; segment I shorter than wide, segment II ing toward R+M (fig. 6), widened antennal seg- slightly longer than wide, its dorsal surface not ment I (as in Namyatova et al., 2016: fig. 8B), elongate posteriorly (as in Namyatova et al., dorsal surface of labial segment II not elongate 2016: fig. 6E); segment III subequal to half of posteriorly (as in Namyatova et al., 2016: fig. 6E), head length ventrally; segment IV ca. 0.3× as right paramere sickle shaped, without outer angle long as segment III. Antenna: Segment widened and with inner angle (fig. 13I), left paramere with (as in Namyatova et al., 2016: fig. 8B), ca. 1.4– flattened apical half, with widened and serrate 1.5× as long as head width, ca. 0.8–0.9× as long outgrowth on inner margin (fig. 13J), ductus sem- as pronotum width; segment II ca. 1.7–1.8× as inis elongate, straight apically (fig. 10L). long as head width, ca. 1.0–1.1× as long as pro- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 139 notum width; segment III slightly longer than notum ca. 1.1× as wide as long and ca. 1.9× as segment II. Thorax: Anterior part of pronotum wide as head. Genitalia: Not dissected. distinctly shorter than posterior part, distinctly Distribution: Ponape Is. (Micronesia) (fig. 20). raised above head; collar not delimited; poste- Host plants: Unknown. rior part slightly upraised; posterior margin Discussion: Felisacus ponaponensis was orig- concave, pronotum ca. 1.0–1.1× as wide as long inally described in the monotypic genus Felisaco- and ca. 1.7–1.8× as wide as head; mesoscutum ris (Carvalho, 1956). Namyatova and Cassis not exposed; inner margin of cuneus convex (as (2016) tentatively placed this genus into the tribe in Namyatova et al., 2016: fig. 13F), its outer Felisacini on the basis of the structural similarity. margin ca. 2.5× as long as base. Abdomen: Gen- Felisacus ponaponensis has the same structural ital capsule rotated left 45° relative to the rest of autapomorphies of Felisacus (see diagnosis of the abdomen. Genitalia: Genital capsule (fig. 15H) genus), it also has an aedeagus very similar to slightly longer than wide; ventral wall ca. 1.5× other species of Felisacus (cf. fig. 10L with figs. as long as dorsal wall, posterior margin of geni- 10C, D, F–I, J, K, O) and the parameres are very tal capsule not curved, right side folded, left similar to those of F. bismarckensis (cf. fig. 11L, side not widened; paramere sockets distinctly M with fig. 13I, J). On this basis we synonymize acute; distance between paramere sockets ca. Felisacoris with Felisacus. Felisacus ponaponensis 0.3× as long as genital capsule width at base; cannot be confused with any other species of paramere sockets distinctly acute; distance Felisacus and we treat it as a distinct species between paramere sockets ca. 0.25× as long as within it, as a new combination. genital capsule width at base. Right paramere Material examined: Paratypes: MICRONE- (fig. 13I) sickle shaped; apex concave; medial SIA: Pohnpei: Mt. Dolennankap, 518 m, 10 Aug part as wide as basal part, without setae, outer 1946, H.K. Townes, 1♂ (00341131) (USNM). margin of medial part convex, inner margin Ponape Is., Matalanium [Matalanim], 6.87835°N concave, without swelling; outer angle absent; 158.22799°E, 11 Aug 1946, R.G Oakley, 1♂ inner angle present, without outgrowth; basal (00338820), 2♀ (00341132, 00338821) (USNM). part of paramere slightly shorter than rest of paramere. Left paramere (fig. 13J) apical part flattened, with toothlike outgrowth apically, Felisacus rubricuneus Carvalho without outgrowth on dorsal surface; inner Figures 6, 9H, 13K, 20 margin with wide, short, serrate outgrowth. Felisacus rubricuneus Carvalho, 1956: 30 (origi- Aedeagus (general view as in Namyatova et al.: nal description). fig. 22M) vesica without spicules; apical half of ductus seminis sclerotized, hooked apically, sec- Diagnosis: Recognized by the following ondary gonopore placed near phallotheca combination of characters: mostly dark color- mouth in repose (fig. 10L). ation; antennal segment I brown to black, anten- Female. Total length 3.6–3.8. COLORATION: nal segment II yellow, head reddish brown; As in male. STRUCTURE AND MEASURE- pronotum brown to black laterally with reddish MENTS: As in male, segment IV ca. 0.3x as long brown stripe medially; marking along inner mar- as segment III. STRUCTURE AND MEASURE- gin of corium brown, wide, not extending toward MENTS: Structure as in male; body ca. 3.4–3.5× R+M anteriorly; cuneus red, cylindrical antennal as long as pronotum width; vertex ca. 2.8–2.9× segment I (as in Namyatova et al., 2016: fig. 8A), as wide as eye; antennal segment I ca. 1.5× as vertex upraised (as in Namyatova et al., 2016: fig. long as head width, ca. 0.8× as long as pronotum 6D); body length in male 3.6–3.8, inner part of width; segment II ca. 1.6–1.7× as long as head cuneus brownish red; medial part of right para- width, ca. 0.9× as long as pronotum width; pro- mere as wide as basal part, shorter than basal 140 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 and apical parts combined, its inner margin wid- segment III. Antenna: Segment I distinctly lon- ened (Carvalho, 1956: fig. 17a), vesica with five ger than head width, cylindrical, ca. 1.5× as long spicules, none of which can be homologized with as head width, ca. 0.9× as long as pronotum the spicules of other species, one of spicules large width; segment II ca. 2.0–2.1 as long as head and suboval (fig. 9H). width, ca. 1.2–1.3× as long as pronotum width. Description: Male. Total length 3.6–3.8. Thorax: Anterior part of pronotum slightly COLORATION (fig. 6): Head: Yellow to pale shorter than posterior part; collar delimited; brown with reddish tinge dorsally and ventrally, posterior part slightly upraised; posterior mar- yellow anteriorly and brown to reddish brown gin of pronotum slightly concave; pronotum ca. laterally. Eye dark brown to black sometimes 1.1–1.2× as wide as long and 1.6–1.7× as wide as with reddish tinge. Labium: Uniformly yellow or head. Hemelytron: Area along inner margin of with segment IV pale brown. Antenna: Segment corium almost flat; inner margin of corium con- I brown with reddish tinge; segment II yellow, vex (as in Namyatova et al., 2016: fig. 13E), ca. segment III yellow basally gradually darkening 3× as long as base. Abdomen: Genital capsule apically. Thorax: Pronotum mostly brown, yel- rotated left at right angle relative to rest of abdo- low to pale brown medially; scutellum pale men. Genitalia: Left paramere (fig. 13K) brown; pleura including scent gland evaporative L-shaped, apical part flattened, with toothlike area whitish yellow. Hemelytron: Mostly translu- outgrowth on posterior side medially (as in fig. cent, colorless; clavus and marking along inner 11G) and without outgrowth on dorsal surface; margin of corium posteriorly brown; cuneus middle part widened, without swelling or out- dark red; membrane with yellow or grayish growth; setae only on middle part near outer tinge, cell yellow with reddish tinge. Legs: Mostly margin. Aedeagus (general view as in Namya- yellow, hind tibia and tarsi dark yellow to pale tova et al., 2016: fig. 22I) conjunctiva weakly brown. Abdomen: Yellow. SURFACE AND VES- sclerotized, sclerotized part of ductus seminis TITURE: Dorsum and antennal segments I and very short, secondary gonopore placed at base of II, with scarce setae mostly shorter than anten- vesica in repose; vesica with five spicules, none nal segment II diameter; femora covered with of which can be homologized with the spicules setae longer than antennal segment II diameter. of other species (fig. 9H). STRUCTURE AND MEASUREMENTS: Body Distribution: Caroline Is. (Kusaie) (fig. 20). ca. 4.3–4.5× as long as pronotum width. Head: Host plants: Unknown. Depression delimiting occipital region present Discussion: Females were not examined. dorsally and laterally (as in Namyatova et al., Due to its coloration F. rubricuneus cannot be 2016: fig. 4E); distance between depression and confused with any other Felisacus species. Super- pronotum shorter than eye diameter; longitudi- ficially it is similar to other dark-colored species nal sulcus on dorsal surface longer than eye with a red cuneus, such as F. bau, F. bellus, F. diameter; distance from eye to pronotum longer malayensis, F. senaru, and F. signis. However, all than eye diameter, slightly swollen laterally (as those species differ in the posterior part of pro- in Namyatova et al., 2016: fig. 4E); vertex 2.0– notum being mostly yellow to pale brown and 2.2× as wide as eye, upraised. Labium (as in the antennal segment II has similar coloration or Namyatova et al., 2016: figs. 6D, 9C): Reaching darker than antennal segment I. middle posterior margin of mesosternum; seg- Material examined: Paratypes: MICRONE- ments I and II combined longer than half of seg- SIA: Kosrae: Mt. Tafeyat, Kusaie Is., 5.3134°N ment III; segment I shorter than wide; segment 162.977°E, 305 m, 21 Aug 1946, H.K. Townes, II slightly longer than wide, elongate dorsally; 1♂ (00338822) (USNM). Mt. Tafeyat, Kusaie Is., segment III slightly shorter than ventral side of 5.3134°N 162.977°E, 198 m, 20 Aug 1946, H.K. head; segment IV more than 2.5–3× as long as Townes, 1♂ (00338823) (USNM). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 141
Felisacus schuhi, sp. nov. area whitish yellow. Hemelytron: Mostly colorless and translucent; inner part of clavus sometimes Figures 6, 9J, 13L, M, 15I, 20 opaque, often brown with small markings and Diagnosis: Recognized by the following margins brown to dark brown, rarely inner part combination of characters: dorsal surface of head of clavus yellow to pale brown with pale brown whitish yellow to pale brown; lateral sides of col- to brown margins with or without small brown lar yellow to pale brown; pronotum mostly yel- markings; outer part of clavus often yellow to low, sometimes with a pair of pale brown pale brown apically; marking along inner margin markings near humeral angles; inner part of cla- of corium pale brown to brown, sometimes paler vus often brown, sometimes yellow to pale posteriorly; embolium with margins pale brown brown; cuneus mostly colorless; cylindrical to brown, sometimes embolium yellow apically antennal segment I (as in Namyatova et al., 2016: or uniformly yellow; cuneus with yellow or pale fig. 8A), vertex upraised (as in Namyatova et al., brown margins, sometimes outer margin and 2016: fig. 6D); labium slightly surpassing middle apex pale brown; membrane sometimes with of mesosternum; antennal segment I ca. 1.5–1.9× grayish tinge, membrane cell yellow. Legs: Coxae as long as head width; cuneus ca. 3× as long as whitish yellow; femora whitish yellow basally base; medial part of right paramere slightly wider and pale brown, sometimes with reddish mark- than basal part, shorter than basal and apical ing, apically; tibiae varying from yellow to pale parts combined; its inner margin convex (fig. brown, sometimes whitish yellow apically, some- 13L); apical part of left paramere not widened, times apical part or entire tibia with reddish ca. 3× as long as wide (fig. 13M); vesica with five tinge; tarsi pale brown to brown, often with seg- spicules, including spicules A, B, and C (fig. 9J). ment I and rarely segment II whitish yellow to Description: Male. Total length 3.8–4.3. yellow. Abdomen: Whitish yellow or yellow, COLORATION (fig. 6): Head: Whitish yellow to sometimes with pale brown to brown dorsal sur- pale brown, usually with red markings, dorsal face of pregenital segments and genital capsule. surface usually darker than lateral and ventral SURFACE AND VESTITURE: Corium smooth, sides; sometimes head uniformly colored. Eye with shallow and scarce punctures. Dorsum dark brown with reddish tinge, rarely pale clothed with setae subequal to or shorter than brown. Labium: Whitish yellow to pale brown, antennal segment II diameter; antennal segment segments III–IV usually darker than segments I and femora clothed with suberect setae shorter I–II and with reddish tinge, sometimes labium than antennal segment II diameter; abdomen uniformly colored. Antenna: Segment I yellow to clothed with short, erect setae. STRUCTURE pale brown, paler basally and sometimes dark- AND MEASUREMENTS: Body ca. 4.2–4.5× as ened apically; segment II pale brown, reddish long as pronotum width. Head: Depression brown or brown, often darker than segment I; delimiting occipital region distinct dorsally and segments III–IV brown to dark brown, often laterally (as in Namyatova et al., 2016: fig. 4E); darker than segment II. Thorax: Pronotum yel- distance between depression and pronotum dis- low to pale brown, with brown anterior margin, tinctly shorter than eye diameter; longitudinal sometimes marking along posterior margin or at sulcus on dorsal surface longer than eye diame- least marking near humeral angles also brown, ter; distance from eye to pronotum distinctly reddish markings laterally sometimes present; longer than eye diameter, not swollen laterally mesoscutum and scutellum whitish yellow to (as in Namyatova et al., 2016: fig. 4E); vertex ca. yellow, sometimes depression and punctures 1.5–1.9× as wide as eye, upraised (as in Namya- between them pale brown; scutellum sometimes tova et al., 2016: fig. 6D). Labium (as in Namya- pale brown to brown apically; thoracic pleura tova et al., 2016: figs. 6D, 9C): Surpassing middle yellow to pale brown; scent gland evaporative of mesosternum but distinctly not reaching pos- 142 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 terior margin of mesosternum; segments I and II 2016: fig. 22I) conjunctiva weakly sclerotized; strongly reduced, combined shorter than half of secondary gonopore placed at base of vesical in segment III; segment I shorter than wide; seg- repose; sclerotization of ductus seminis around ment II slightly longer than wide, its dorsal sur- secondary gonopore shorter than wide; vesical face elongate posteriorly; segment III subequal with five spicules, including large spicules A, B, to ventral side of head; segment IV ca. 1.5× as and C (fig. 9J). long as segment III. Antenna: Segment I cylin- Female. Total length 4.3–4.6. COLORATION drical (as in Namyatova et al., 2016: fig. 8A), ca. (fig. 6): Head: Similar to male, rarely uniformly 1.5–1.9× as long as head width, ca. 1.0–1.2× as pale brown. Labium: Similar to male, sometimes long as pronotum width; segment II ca. 1.8–2.2× uniformly pale brown. Antenna: Similar to male, as long as head width, ca. 1.3–1.5× as long as antennal segment I rarely uniformly pale brown. pronotum width; segment III slightly longer Thorax: As in male. Legs and hemelytron: As in than segment II; segment IV ca. 0.3× as long as male. Abdomen: Yellowish to pale brown, often antennal segment III. Thorax: Anterior part of with reddish apex and brownish dorsal surface. pronotum slightly shorter than posterior part; SURFACE AND VESTITURE: As in male. collar delimited; posterior part of pronotum STRUCTURE AND MEASUREMENTS: Struc- slightly upraised; posterior margin of pronotum ture as in male; body ca. 4.2–4.5× as long as pro- concave, pronotum ca. 1.2–1.4× as wide as long notum width; vertex ca. 1.6–2.1× as wide as eye; and ca. 1.4–1.5× as wide as head; mesoscutum antennal segment I ca. 1.5–1.8× as long as head exposed. Hemelytron: Area along inner margin width, ca. 1.0–1.1× as long as pronotum width; of corium almost flat; inner margin of cuneus segment II ca. 1.8–2.2× as long as head width, ca. convex (as in Namyatova et al., 2016: fig. 13E), 1.2–1.4× as long as pronotum width; pronotum outer margin of cuneus ca. 3× as long as base. ca. 1.2–1.3× as wide as long and ca. 1.5–1.7× as Abdomen: Genital capsule rotated left at right wide as head. Genitalia (as in Namyatova et al., angle relative to rest of abdomen. Genitalia: 2016: figs 23F, G): Dorsal labiate plate wider than Genital capsule (fig. 15I) ca. 1.5× as long as wide; distance between apodemes of second valvula; ventral wall ca. 1.7× as long as dorsal wall, with mostly smooth, without distinct striations or lon- posterior margin of ventral wall semioval, gitudinal membranous ridge, with semicircular smooth, without outgrowth(s), its apex slightly sclerites and distinct sclerotized rings laterally; shifted left, not curved; sides of genital capsule lateral oviducts placed almost medially, very close not modified; right paramere socket acute, left to each other, spermathecal gland placed between socket rounded; distance between paramere lateral oviducts; dorsal labiate plate with distinct sockets subequal to half of genital capsule width tubercles, without membranous lobe medially. at base. Right paramere (fig. 13L) distinctly Distribution: Loyalty Is., Solomon Is. (Cho- curved in apical half; apex straight; medial part iseul Is., Santa Cruz), Vanuatu (Malekula Is., Epi slightly wider than basal part, bearing setae, with Is., Santo Is.) (fig. 20). outer margin straight or slightly concave and Host plants: Unknown. inner margin slightly widened; outer angle dis- Etymology: The species is named after Ran- tinct, not swollen; inner angle rounded, without dall Schuh, emeritus curator of the American setae; basal part ca. 0.15–0.2× as long as rest of Museum of Natural History, who kindly loaned paramere. Left paramere (fig. 13M) L-shaped; us specimens of Felisacus. apical part not flattened, with tooth on posterior Discussion: Felisacus schuhi is similar to F. side medially and without outgrowth on dorsal elegantulus, F. lordhowensis, and F. meilingae in surface; middle part widened, without swelling coloration and structure (figs. 5, 6). Felisacus ele- or outgrowth; setae only on middle part near gantulus differs from F. schuhi in the humeral outer margin. Aedeagus (as in Namyatova et al., angles often brown to dark brown, the inner part 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 143 of the clavus mostly colorless or yellow and reach- Felisacus senaru, sp. nov. ing posterior margin of the mesosternum; the Figures 6, 9I, 13N, O, 20 vesica has a short spicule B and lacks spicules A Diagnosis: Recognized by the following com- and C, but with a large bifurcate spicule (fig. 8H). bination of characters: relative dark coloration; Felisacus lordhowensis is separated from F. schuhi head yellow to pale brown; posterior part of pro- by the inner part of the clavus whitish yellow to notum yellow to pale brown with wide brown yellow, the labium reaching or slightly surpassing stripe along posterior margin; antennal segment I the posterior margin of the metasternum, and the mostly brown and segment II reddish brown; vesica has six spicules, with a short spicule B and without spicules A and C (fig. 8R). Felisacus mei- antennal segment I cylindrical (as in Namyatova lingae differs from F. schuhi by the posterior mar- et al., 2016: fig. 8A); vertex upraised (Namyatova gin of the pronotum with a brown marking et al., 2016: fig. 6D); body length in male 3.6–3.8, posteriorly and the presence of six vesical spicules, in female 3.8–4.1; medial part of right paramere including spicules A, B, and C (fig. 9J). slightly wider than basal part, with outer margin Material examined: Holotype: VANUATU: concave (fig. 13N), vesical with six spicules, Shefa: Epi Island: Lowekewou, 16.6°S 168.1667°E, including ling spicules A, B, and C, spicule C dis- 10 m, 31 Aug 1979, B.H. Gagne, 1♂ (00043026) tinctly moved posteriorly (fig. 9I). (BPBM). Paratypes: EAST TIMOR: Estacao Zoo- Description: Male. Total length 3.6–3.8. tecnica and foot of Mundo Perdido nr Ossu, COLORATION (fig. 6): Head: Yellow to pale Timor Portugues, 8.73423°S 126.35914°E, 09 Mar brown with reddish tinge dorsally. Eye dark 1961, G.F. Gross, 1♂ (00017812) (SAMA). NEW brown to black. Labium: Yellow. Antenna: Seg- CALEDONIA: Province des Iles Loyauté: Fay- ment I brown, pale yellow to yellow basally and aoue, Ouvea Is., 20.65774°S 166.54238°E, 25 m, ventrally; segment II reddish brown; segments Dec 1968, N.L.H. Krauss, 2♂ (00042305, III–IV dark brown to black, darker than segment 00042307), 1♀ (00042308) (BPBM). SOLOMON II. Thorax: Pronotum yellow to pale brown, with ISLANDS: Isabel: Sasamungga [Sasamonga], brown anterior margin and wide brown stripe 7.03333°S 156.78333°E, 50 m, Feb 1984, N.L.H. along posterior margin; mesoscutum pale Krauss, 1♂ (00043242) (BPBM). Makira-Ulawa: brown, scutellum brown; thoracic pleura yellow; Graciosa Bay, 10.75°S 165.76666°E, 25 m, Jan scent gland evaporative area whitish yellow. 1977, N.L.H. Krauss, 1♂ (00043241), 1♀ Hemelytron: Mostly colorless and translucent; (00043243) (BPBM). VANUATU: Espiritu Santo inner part of clavus translucent or opaque, Is.: Big Bay, E Jordan River, 15.17443°S brown; marking along inner margin of corium 166.92759°E, 20 m, 12 Sep 1979, B.H. Gagne, 2♂ brown; embolium with margins yellow; cuneus (00042314, 00042315), 4♀ (00043043–00043046) with pale brown margins and inner half red; (BPBM). Malampa: South West Bay, Malekula Is., membrane with greyish tinge. Legs: Coxae whit- 16.4722°S 167.43124°E, 13 Oct 1971, P. Cochereau, ish yellow; femora, tibiae, and tarsi yellow. Abdo- 1♂ (00017553), 3♀ (00017554–00017556) men: Yellow with greenish tinge. SURFACE (SAMA). Penama: Maewo Is.: Sounwari, 15.38°S AND VESTITURE: Corium smooth, impunc- 168.12989°E, 50 m, 04 Sep 1979–05 Sep 1979, tate. Dorsum clothed with setae suberect and B.H. Gagne, 1♀ (00043062) (BPBM). Shefa: Epi longer than antennal segment II diameter; Island: Lowekewou, 16.6°S 168.1667°E, 10 m, 31 antennal segment I and femora clothed with Aug 1979, B.H. Gagne, 4♂ (00043023–00043025, suberect setae shorter than antennal segment II 00043027), 15♀ (00043028–00043042) (BPBM). diameter; abdomen clothed with short erect Vaemali, 16.58959°S 168.16855°E, 125 m, 06 Aug setae. STRUCTURE AND MEASUREMENTS: 1967–10 Aug 1967, J. and M. Sedlacek, 1♂ Body ca. 4.4–4.7× as long as pronotum width. (00042313) (BPBM). Head: Depression delimiting occipital region 144 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 present dorsally and laterally (as in Namyatova out outgrowth on dorsal surface; middle part et al., 2016: fig. 4E); distance between depression widened, without swelling or outgrowth; setae and pronotum shorter than eye diameter; longi- present only on middle part near outer margin. tudinal sulcus on dorsal surface longer than eye Aedeagus (general view as in Namyatova et al., diameter; distance from eye to pronotum dis- 2016: fig. 22I) conjunctiva weakly sclerotized; tinctly longer than eye diameter, not swollen lat- secondary gonopore placed at base of vesical in erally (as in Namyatova et al., 2016: fig. 4E); repose; sclerotization of ductus seminis around vertex ca. 1.5–1.9× as wide as eye; upraised view secondary gonopore shorter than wide; vesica (Namyatova et al., 2016: fig. 6D). Labium (as in with six spicules, including spicules A, B, and C, Namyatova et al., 2016: figs. 6D, 9C): Surpassing spicule C convolute basally and distinctly moved metasternum, but distinctly not reaching poste- posteriorly (fig. 9I). rior margin of metasternum; segments I and II Female. Total length 3.8–4.1. COLORATION strongly reduced, combined shorter than half of (fig. 6): Similar to male, sometimes dorsal side of segment III; segment I shorter than wide; seg- head and pronotum with red or brown tinge, ment II slightly longer than wide, its dorsal sur- antennal segment I sometimes almost uniformly face elongate posteriorly; segment III slightly yellow to pale brown or yellow with reddish tinge; shorter than ventral side of head; segment IV cuneus sometimes yellow with reddish brown mar- twice as long as segment III. Antenna: Segment gins and inner parts, tibiae sometimes brown. SUR- I cylindrical (as in Namyatova et al., 2016: fig. FACE AND VESTITURE: As in male. 8A), ca. 1.7–1.9× as long as head width, ca. 1.1– STRUCTURE AND MEASUREMENTS: Structure 1.2× as long as pronotum width; segment II ca. as in male; body ca. 4.4–4.5× as long as pronotum 2.1–2.3× as long as head width, ca. 1.3–1.5× as width; vertex ca. 1.8–2.5× as wide as eye; antennal long as pronotum width; segment III slightly segment I ca. 1.6–1.8× as long as head width, ca. longer than segment II; segment IV ca. 0.2× as 1.0× as long as pronotum width; segment II ca. long as antennal segment III. Thorax: Anterior 2.0–2.3× as long as head width, ca. 1.2–1.4× as long part of pronotum shorter than posterior part; as pronotum width; pronotum ca. 1.2× as wide as collar delimited; posterior part only slightly long and ca. 1.6–1.7× as wide as head. upraised; posterior margin of pronotum con- Distribution: Specimens were collected only cave, pronotum ca. 1.2× as wide as long and ca. in Senaru (Lombok Is.) (fig. 20). 1.6× as wide as head; mesoscutum slightly Etymology: The species is named after the exposed. Hemelytron: Area along inner margin place Senaru (Lombok, Indonesia), where it was of corium flat; inner margin of cuneus concave collected. (as in Namyatova et al., 2016: fig. 13E), outer Host plants: Plesioneuron sp. (Thelypterida- margin of cuneus ca. 2.5× as long as base. Abdo- ceae), many specimens. men: Genital capsule rotated left at right angle Discussion: Felisacus senaru can be confused relative to rest of abdomen. Genitalia: Right with other dark specimens of this genus that pos- paramere (fig. 13N) apical part distinctly delim- sess a red or partially red cuneus, such as F. bau, ited, only slightly concave posteriorly, medial F. bellus, F. jacobsoni, F. malayensis, and F. signis. part slightly wider than basal part, bearing setae, These species differs from F. senaru in the cuneus with outer margin straight or slightly concave entirely or mostly red and in the shape and/or and inner margin slightly widened; outer angle number of vesical spicules (cf. fig. 9I with figs. distinct, not swollen; inner angle rounded, with- 8D, E, M, 9A. K). out setae; basal part ca. 0.2× as long as rest of Material examined: Holotype: INDONE- paramere. Left paramere (fig. 13O) L-shaped; SIA: West Nusa Tenggara: Lombok Island, nr apical part not flattened, ca. 3× as long as wide, Senaru, 8.31689°S 116.40381°E, 29 Aug 2012, F. with tooth on posterior side medially and with- Konstantinov, 1♂ (00386472) (ZISP). Paratypes: 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 145
INDONESIA: West Nusa Tenggara: Lombok brown, with reddish tinge, ventral side yellow to Island, Senaru, Panorama track to Sendang Gila pale brown. Eye dark brown to black. Labium: Waterfall, 8.30111°S 116.40833°E, 28 Aug 2012, Yellow. Antenna: Segment I yellow with reddish F. Konstantinov, 1♂ (00386469) (ZISP); 30 Aug tinge, segment II reddish brown; segment III 2012, F. Konstantinov, Plesioneuron sp. (Thelyp- brown. Thorax: Pronotum and scutellum yellow teridaceae), det. Michael Lovave (LAE herbar- to pale brown; humeral angle with pale brown to ium, PNG), 12♂ (00386488–00386499), 1 sex brown marking, anterior margin sometimes pale unknown (00386500), 12♀ (00386501– brown; thoracic pleura yellow; scent gland evap- 00386512) (ZISP). Lombok Island, Senaru, track orative area whitish yellow. Hemelytron: Mostly to Tiu Kelep Waterfall, 8.30111°S 116.40833°E, translucent, colorless; inner part of clavus with 28 Aug 2012, F. Konstantinov, 2♀ (00386461, yellow tinge and reddish yellow margins; corium 00386462) Plesioneuron sp. (Thelypteridaceae), with brown marking along apical part of clavus, 4♀ (00386463–00386466) (ZISP); 31 Aug 2012, area along inner margin of corium pale brown to F. Konstantinov, Plesioneuron sp. (Thelypterida- brown; embolium with pale brown or red mar- ceae), 1 sex unknown (00386468) (ZISP). Lom- gins; cuneus red. Legs: Whitish yellow to yellow, bok Island, nr Senaru, 8.31689°S 116.40381°E, 29 sometimes femora and tibiae with reddish tinge. Aug 2012, F. Konstantinov, 13♀ (00386474– Abdomen: Yellow to pale brown. SURFACE AND 00386480, 00386474–00386479), 1 sex unknown VESTITURE: Dorsum, antennal segment I and (00386480), 2♂ (00386470, 00386471) (ZISP). II and femora with setae shorter than antennal Lombok Island, nr Senaru, 8.31956°S 116.405°E, segment II diameter; abdomen clothed with 31 Aug 2012, F. Konstantinov, Plesioneuron sp. short setae. STRUCTURE AND MEASURE- (Thelypteridaceae), det. Michael Lovave (LAE MENTS: Body ca. 4.3× as long as pronotum herbarium, PNG), 4♂ (00386481–00386484), width. Head: Depression delimiting occipital 3♀ (00386485–00386487) (ZISP). region present dorsally and laterally (as in Namy- atova et al., 2016: fig. 4E); distance between depression and pronotum distinctly shorter than Felisacus signis Hsiao, status nov. eye diameter; longitudinal sulcus on dorsal sur- Figures 6, 9K, 13P, Q, 20 face subequal to eye diameter; distance from eye Felisacus carpenterae signis Hsiao, 1944: 386. to pronotum subequal to or slightly longer than eye diameter, not swollen laterally (as in Namya- Diagnosis: Recognized by the following combi- tova et al., 2016: fig. 4E); vertex ca. 1.4× as wide nation of characters: head and pronotum yellow to as eye, upraised (as in Namyatova et al., 2016: fig. pale brown; posterior part of pronotum with dark 6D). Labium (as in Namyatova et al., 2016: figs. brown to black markings at humeral angles; anten- 6D, 9C): Segments I and II strongly reduced, nal segment I yellow with reddish tinge; cuneus combined subequal to half of segment III; seg- red; marking along inner margin of corium brown, ment I shorter than wide; segment II almost as not extending toward R+M anteriorly; cylindrical long as wide, elongate dorsally; segment III antennal segment I (as in Namyatova et al., 2016: slightly shorter than ventral side of head; seg- fig. 8A), vertex upraised (as in Namyatova et al., ment IV twice as long as segment III. Antenna: 2016: fig. 6D); length of body in male 3.3, length in Segment I distinctly longer than head width, female 3.35; right paramere with medial part cylindrical (as in Namyatova et al., 2016: fig. 8A), shorter than apical part and distinctly wider than ca. 1.6× as long as head width, ca. 1.2× as long as basal part (fig. 13P); vesica with five spicules, pronotum width; segment II ca. 2× as long as including spicule B (fig. 9K). head width, ca. 1.4× as long as pronotum width. Description: Male. Total length 3.3. COL- Thorax: Anterior part of pronotum distinctly ORATION (fig. 6): Head: Mostly yellow to pale shorter than posterior part; collar distinct; pos- 146 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 terior part only slightly swollen; lateral margins junior synonym of F. javanus (see discussion for not delimited with depression; posterior margin this species), and F. carpenterae signis differs in of pronotum straight; pronotum ca. 1.2× as wide coloration and male genitalia. On this basis we as long and 1.4× as wide as head. Hemelytron: decided to raise this taxon to species rank. Area along inner margin of corium almost flat; Felisacus signis differs from F. javanus in having inner margin of cuneus convex (as in Namyatova darker coloration, the humeral angles are dark et al., 2016: fig. 13E), outer margin of cuneus ca. brown to black and the cuneus is or entirely red. 2.5× as long as base. Abdomen: Genital capsule Felisacus javanus also differs in the shape of the rotated left at right angle relative rest of abdo- right paramere, with the medial part longer than men. Genitalia: Right paramere (fig. 13P) apical the apical part and almost as wide as basal part (cf. part distinct; apex slightly concave; medial part fig. 12G with fig. 13P), and the presence of the distinctly wider than basal part, bearing setae, vesical spicules A, B, and C (cf. fig. 8N with fig. with outer margin straight slightly concave and 9K). Felisacus signis is similar to other dark species inner margin swollen; outer angle distinct, of this genus with a red cuneus, including F. bau, slightly swollen; inner angle absent, without F. bellus, F. jacobsoni, F. malayensis, and F. senaru. setae; basal part ca. 0.25× as long as rest of para- However, all those species differ from F. signis in mere. Left paramere (fig. 13Q) L-shaped; apical shape and/or number of vesical spicules (cf. fig. part not flattened, ca. 3× as long as wide, with 9K with figs. 8D, E, M, 9A, I). toothlike outgrowth on posterior side medially Material examined: Paratypes: MALAY- and without outgrowth on dorsal surface; middle SIA: Sabah: Sandakan, Borneo, 5.84561°N part widened, without swelling or outgrowth(s); 118.05705°E, 66 m, no date provided, Baker, 1♂ setae only on middle part near outer margin. (00338827), 1♀ (00338828) (USNM). Aedeagus (general view as fig. 22I in Namyatova et al., 2016) conjunctiva weakly sclerotized; sec- ondary gonopore placed at base of vesical in Felisacus solomonicus, sp. nov. repose; sclerotization of ductus seminis around Figures 7, 9L, 13R, S, 15J, 20 secondary gonopore shorter than wide; vesica Diagnosis: Recognized by the following with five spicules, including elongate spicule B, combination of characters: head yellow to pale not swollen medially (fig. 9K). brown; pronotum mostly yellow, antennal seg- Female. Total length 3.5. COLORATION (fig. ment I red or brown; outer part of clavus yel- 6): As in male. Abdomen: Yellow, sometimes with low; cuneus yellow with reddish tinge, reddish markings. SURFACE AND VESTITURE: sometimes outer part almost colorless; cylindri- As in male. STRUCTURE AND MEASURE- cal antennal segment I (as in Namyatova et al., MENTS: Body ca. 4.2× as long as pronotum 2016: fig. 8A), vertex upraised (as in Namyatova width; vertex ca. 1.4× as wide as eye; antennal et al., 2016: fig. 6D); labium slightly surpassing segment I ca. 1.6× as long as head width, ca. 1.1× middle of mesosternum; cuneus ca. 3× as long as long as pronotum width; segment II ca. 2.0× as base; medial part of right paramere as wide as long as head width, ca. 1.3× as long as prono- as basal part, shorter than basal and apical parts tum width; pronotum ca. 1.2× as wide as long combined, its outer and inner margins straight, and ca. 1.5× as wide as head. apical part of right paramere distinctly concave Distribution: Singapore (fig. 20). (fig. 13R); vesica with six spicules, including Host plants: Unknown. spicule A (fig. 9L). Discussion: Felisacus signis was originally Description: Male. Total length 3.1–3.4. described as a subspecies of F. carpenterae COLORATION (fig. 7): Head: Mostly yellow to (Hsiao, 1944). Examination of the type material pale brown; tubercle around antennal fossa whit- from USNM showed that F. carpenterae is a ish yellow; longitudinal sulcus and buccula 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 147 sometimes pale brown, slightly darker than rest tova et al., 2016: fig. 6D). Labium (as in Namya- of head, and clypeus rarely reddish apically. Eyes tova et al., 2016: figs. 6D, 9C): Surpassing middle brown with reddish tinge. Labium: Yellow, seg- of mesosternum; segments I and II strongly ment III reddish ventrally, segments I–III some- reduced, combined subequal to half of segment times with reddish tinge. Antenna: Segment I III; segment I shorter than wide; segment II reddish brown to brown with yellow base, seg- slightly longer than wide, its dorsal surface elon- ments II–IV brown. Thorax: Yellow with anterior gate posteriorly; segment III slightly longer than margin pale brown and posterior angles reddish ventral side of head; segment IV ca. 1.5× as long or brown; scutellum whitish yellow to yellow, as segment III. Antenna: Segment I cylindrical mesoscutum and scutellum yellow to yellowish (as in Namyatova et al., 2016: fig. 8A), ca. 1.1– brown, punctures between mesoscutum and scu- 1.4× as long as head width, ca. 0.9–1.1× as long tellum pale brown; thoracic pleura yellow, scent as pronotum width; segment II ca. 1.8–1.9× as gland evaporative area yellow to pale brown long as head width, ca. 1.4× as long as pronotum often with whitish yellow apex. Hemelytron: width; segment III slightly longer than segment Mostly translucent, colorless; inner part of clavus II. Thorax: Anterior and posterior parts subequal translucent or opaque, yellow to pale brown, in length; calli delimited; posterior part slightly margins sometimes somewhat darker; outer part upraised; posterior margin not delimited with of clavus sometimes yellow; area along inner depression; posterior margin concave, pronotum margin of corium yellow or pale brown; embo- ca. 1.1–1.2× as wide as long and ca. 1.3–1.4× as lium yellow apically and pale brown to brown wide as head; mesoscutum exposed. Hemelytron: margins; cuneus yellow with reddish tinge, Area along inner margin of corium almost flat; sometimes outer part almost colorless, margin inner margin of cuneus convex (as in Namyatova brown or reddish; membrane sometimes with et al., 2016: fig. 13E), outer margin ca. 3× as long grayish tinge, membrane call yellow to pale as base. Abdomen: Genital capsule rotated 45° brown. Legs: Coxae whitish yellow, femora whit- relative to rest of abdomen. Genitalia: Genital ish yellow basally and yellow apically with red capsule (fig. 15J) ca. 1.5× as long as wide; ventral marking on apex; tibiae reddish; tarsi yellow to wall ca. 1.5× as long as dorsal wall, its posterior pale brown. Abdomen: Reddish, yellow apically. margin smooth, semioval, without outgrowth(s), SURFACE AND VESTITURE: Corium smooth, not curved dorsally; its apex slightly shifted often with shallow and scarce punctures. Dor- right; right side of genital capsule not curved, left sum with suberect setae subequal to or shorter side not widened; paramere sockets more or less than antennal segment II diameter; antennal seg- rounded; distance between paramere sockets ca. ment II and femora with suberect setae shorter 0.7× as long as genital capsule width at base. than antennal segment II diameter; abdomen Right paramere (fig. 13R) distinctly curved in clothed with suberect mostly short simple setae. apical half; apex concave posteriorly; medial part STRUCTURE AND MEASUREMENTS: Body wider than basal part, bearing setae, with outer ca. 4.3–4.8× as long as pronotum width. Head: margin straight and inner margin slightly con- Depression delimiting occipital region present cave, without swelling; outer angle distinct; inner dorsally and laterally (as in Namyatova et al., angle indistinct, without setae; basal part of para- 2016: fig. 4E); distance between depression and mere ca. 0.15–0.2× as long as rest of paramere. pronotum distinctly shorter than eye diameter; Left paramere (fig. 13S) L-shaped; apical part not longitudinal sulcus on dorsal surface longer than flattened, with tooth on posterior side medially eye diameter; distance from eye to pronotum (as in fig. 11G) and without outgrowth on dorsal longer than eye diameter, not swollen laterally surface; middle part widened, without swelling (as in Namyatova et al., 2016: fig. 4E); vertex ca. or outgrowth; setae only on middle part near 1.6–1.8× as wide as eye, upraised (as in Namya- outer margin. Aedeagus (as in Namyatova et al., 148 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
2016: fig. 22I) conjunctiva weakly sclerotized; (BPBM). Paratypes: SOLOMON ISLANDS: secondary gonopore placed at base of vesical in Malaita: Auki, 8.767°S 160.7°E, 100 m, Feb 1985, repose; sclerotization of ductus seminis around N.L.H. Krauss, 1♀ (00043233) (BPBM). Tagata- secondary gonopore shorter than wide; vesica lau [Tangtalau], 8.73333°S 160.73333°E, 200 m, with six spicules, including elongate and narrow 25 Sep 1957, J.L. Gressitt, 1♂ (00043229) spicule A (fig. 9L). (BPBM). New Georgia Islands: Gizo, Gizo Is., Female. Total length 3.3–3.4. COLORATION 8.1°S 156.85°E, 100 m, Nov 1975, N.L.H. Krauss, (fig. 7): Similar to male, thorax yellow with ante- 2♂ (00043227, 00043228) (BPBM). Tamaneke, rior margin pale brown and posterior angles red- 8.48333°S 158.06667°E, 50 m, Feb 1987, N.L.H. dish, evaporatorium uniformly pale brown; Krauss, 1♀ (00043232) (BPBM). Western Prov- abdomen uniformly reddish or pale brown api- ince: New Georgia Islands: Munda, New Geor- cally. SURFACE AND VESTITURE: As in male. gia Is., 8.31666°S 157.25°E, 50 m, Feb 1984, STRUCTURE AND MEASUREMENTS: Struc- N.L.H. Krauss, 1♂ (00043230) (BPBM). ture as in male; body ca. 4.2–4.4× as long as pro- notum width; vertex ca. 1.7–1.9× as wide as eye; Felisacus tanna, sp. nov. antennal segment I ca. 1.3–1.4× as long as head width, ca. 0.9–1.0× as long as pronotum width; Figures 7, 9M, 13T, U, 15K, 20 segment II ca. 1.7–1.8× as long as head width, ca. Diagnosis: Recognized by the following 1.2–1.3× as long as pronotum width; pronotum combination of characters: head yellow to pale ca. 1.1–1.2× as wide as long and ca. 1.4× as wide brown dorsally; pronotum, including collar and as head. Genitalia (as in Namyatova et al., 2016: humeral angles, mostly whitish yellow to yellow; fig. 23F, G): Dorsal labiate plate wider than dis- inner margin of clavus yellow; scent gland evap- tance between apodemes of second valvula; orative area whitish basally and yellow or red mostly smooth, without distinct striations, with apically, rarely uniformly red; cuneus colorless, semicircular sclerite and distinct sclerotized often with yellow tinge, with margins yellow to rings laterally; lateral oviducts placed almost pale brown; cylindrical antennal segment I (as in medially, very close to each other, spermathecal Namyatova et al., 2016: fig. 8A), vertex upraised gland placed between lateral oviducts; dorsal (as in Namyatova et al., 2016: fig. 6D); labium labiate plate with distinct tubercles, without reaching posterior margin of metasternum or membranous lobe medially. slightly surpassing it; setae on femora distinctly Remarks: Antennal segment IV lost in males. shorter than antennal segment II width; medial Distribution: Solomon Is. (Gizo Is., Malaito part of right paramere distinctly wider than basal Is., Georgia Is., Auki Is.) (fig. 20). part; shorter than basal and apical parts com- Host plants: Unknown. bined; its outer margin straight (fig. 13T); apical Etymology: The species is named after Solo- part of left paramere not widened, ca. 3× as long mon Islands, where it was collected. as wide (fig. 13U); vesica with six spicules, Discussion: Felisacus solomonicus is most including spicules A, B, and E (fig. 9M). similar to F. albus, F. amboinae, F. javanus, and F. Description: Male. Total length 3.4–3.6. linae externally (figs. 4, 5). All these species can COLORATION (fig. 7): Head: Mostly whitish be separated from each other by the apical part yellow to yellow, dorsal surface sometimes with of the right paramere straight or slightly concave reddish tinge and reddish markings. Eye dark (figs. 11A, C, 12G, K) and the shape and/or num- brown, often with reddish tinge. Labium: Whit- ber of the vesical spicules (figs. 8A, B, N, P). ish yellow to yellow, labial segment III with red- Material examined: Holotype: SOLOMON dish stripe ventrally, segment IV often somewhat ISLANDS: Malaita: Auki, 8.767°S 160.7°E, 50 m, darker than segments I–III. Antenna: Segment I Dec 1975, N.L.H. Krauss, 1♂ (00043231) yellow to pale brown often with reddish tinge; 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 149 segment II yellow to brown, sometimes with red- upraised (as in Namyatova et al., 2016: fig. 6D). dish tinge, often darker than segment I; segments Labium (as in Namyatova et al., 2016: figs. 6D, III–IV brown to dark brown often darker than 9C): Reaching posterior margin of metasternum segment II. Thorax: Pronotum whitish yellow to or slightly surpassing it; segments I and II yellow, often with reddish tinge anteriorly or strongly reduced, combined shorter than half of with reddish stripe laterally, collar with brown segment III; segment I shorter than wide; seg- anterior margin; mesoscutum and scutellum ment II slightly longer than wide, its dorsal sur- whitish yellow to yellow; thoracic pleura yellow, face elongate posteriorly; segment III longer than sometimes with reddish tinge; scent gland evap- ventral side of head; segment IV ca. 1.5× as long orative area whitish basally and yellow or red as segment III. Antenna: Segment I cylindrical apically, rarely uniformly red. Hemelytron: (as in Namyatova et al., 2016: fig. 8A), ca. 1.5– Mostly translucent and colorless; inner part of 1.8× as long as head width, ca. 0.9–1.1× as long clavus opaque or translucent, yellow, with mar- as pronotum width; segment II ca. 2.0–2.2× as gins yellow to pale brown; marking along inner long as head width, ca. 1.2–1.4× as long as pro- margin of corium yellow; embolium with pale notum width; antennal segment III slightly lon- brown margins and yellow apex; cuneus often ger than segment II; segment IV ca. 0.3× as long with yellow tinge, with margins yellow to pale as segment III. Thorax: Anterior part of prono- brown; membrane with yellow tinge, membrane tum slightly shorter than posterior part; collar cell yellow. Legs: Coxa whitish yellow; femora delimited; posterior part of pronotum slightly whitish yellow basally and yellow to pale brown upraised; posterior angles of pronotum not apically; tibia yellow to pale brown, femora and delimited with depression; posterior margin of tibia sometimes with reddish tinge; tarsi often pronotum straight or concave, pronotum ca. pale brown with segment I paler, yellow to pale 1.3× as wide as long and ca. 1.5–1.7× as wide as brown, sometimes uniformly yellow to pale head; mesoscutum not exposed or exposed. brown. Abdomen: Ventral and lateral sides whit- Hemelytron: Area along inner margin of corium ish yellow to yellow, dorsal surface reddish, geni- flat; inner margin of cuneus convex (as in tal capsule sometimes darker than pregenital Namyatova et al., 2016: fig. 13E), outer margin of segments, yellow to pale brown, rarely abdomen cuneus ca. 3× as long as base. Abdomen: Genital uniformly red. SURFACE AND VESTITURE: capsule rotated left at right angle relative to the Corium smooth, with shallow and scarce punc- rest of abdomen. Genitalia: Genital capsule (fig. tures. Dorsum clothed with setae subequal to or 15K) ca. 1.5× as long as wide; ventral wall ca. shorter than antennal segment II diameter; 1.5× as long as dorsal wall, with posterior margin antennal segment II, femora and abdomen of ventral wall semioval, smooth, without out- clothed with suberect setae distinctly shorter growth, apex inclined leftward, not curved; sides than antennal segment II diameter. STRUC- of genital capsule not modified; paramere socket TURE AND MEASUREMENTS: Body ca. 4.4– slightly acute, left paramere socket rounded; dis- 4.6× as long as pronotum width. Head: tance between paramere sockets subequal to half Depression delimiting occipital region present of genital capsule width at base. Right paramere dorsally and laterally (as in Namyatova et al., (fig. 13T) distinctly curved in apical half; apex 2016: fig. 4E); distance between depression and straight posteriorly; medial part twice as wide as pronotum distinctly shorter than eye diameter; basal part, bearing setae, with outer margin longitudinal sulcus on dorsal surface of head straight and inner margin widened; outer angle slightly longer than eye diameter; distance from distinct, not widened; inner angle rounded, with- eye to pronotum longer than eye diameter, not out setae; basal part of paramere ca. 0.15–0.2× as swollen laterally (as in Namyatova et al., 2016: long as rest of paramere. Left paramere (fig. 13U) fig. 4E); vertex ca. 1.6–2.0× as wide as eye, L-shaped; apical part not flattened, with tooth- 150 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 like outgrowth on posterior side medially (as in medial part of the right paramere (cf. fig. 11AF fig. 11G) and without outgrowth on dorsal sur- with fig. 13T), but the latter species differs by face; middle part widened, without swelling or the labium reaching the middle of the meso- outgrowth; setae only on middle part near outer sternum, the medial part of the right paramere margin. Aedeagus (general view as in Namyatova has the outer margin concave (fig. 11AF) and et al., 2016: fig. 22I) conjunctiva weakly sclero- the vesica has five spicules, including spicules B tized, secondary gonopore placed at base of vesi- and F (fig. 8J). cal in repose; sclerotization of ductus seminis Felisacus tanna can also be confused with F. around secondary gonopore shorter than wide; filicicola, F. lordhowensis, and F. schuhi (figs. 5, 6), vesica with six spicules, including spicules A, B, but these latter three species differ in the middle and E, spicule A short and spicule B large and part of the right paramere being only slightly serrate apically (fig. 9M). wider than the basal part (figs. 11AD, 12S, 13L) Female. Total length 3.5–3.8. COLORATION and the shape and/or number of the vesical spic- (fig. 7): Head and labium: As in male. Antenna: ules (figs. 8I, R, 9J). Similar to male, antennal segment II rarely yel- Material examined: Holotype: VANUATU: low with reddish tinge. Thorax: Similar to male, Taffeta: Tanna Is.: 2 km S Ipeukeul, 19.51511°S but posterior part of pronotum rarely pale brown 169.45466°E, 350 m, 23 Aug 1979, G.A. Samuel- posteriorly. Abdomen: Whitish yellow to yellow son, 1♂ (00043068) (BPBM). Paratypes: VANU- with red dorsal surface. SURFACE AND VESTI- ATU: Penama: Maewo Is.: Sounwari, 15.38°S TURE: As in male. STRUCTURE AND MEA- 168.12989°E, 50 m, 04 Sep 1979–05 Sep 1979, SUREMENTS: Structure as in male; body ca. B.H. Gagne, 3♂ (00043052–00043054), 2♀ 4.2–4.7× as long as pronotum width; vertex ca. (00043061, 00043063) (BPBM). Taffeta: Tanna 1.8–2.2× as wide as eye; antennal segment I ca. Is.: 2 km S Ipeukeul, 19.51511°S 169.45466°E, 1.6–1.7× as long as head width, ca. 1.0–1.1× as 350 m, 23 Aug 1979, G.A. Samuelson, 11♂ long as pronotum width; segment II ca. 1.9–2.1× (00043069–00043072, 00043077–00043083), 3 as long as head width, ca. 1.1–1.3× as long as juvenile (00043073–00043075), 1 sex unknown pronotum width; pronotum ca. 1.2–1.4× as wide (00043066), 1♀ (00043076) (BPBM). Loounap- as long and ca. 1.5–1.7× as wide as head. Genita- kaukangeus, 19.5°S 169.3°E, 350 m, 03 Mar 1970, lia (as in Namyatova et al., 2016: fig. 22I): Dorsal N.L.H. Krauss, 1♂ (00043067) (BPBM). labiate plate wider than distance between apodemes of second valvula; mostly smooth, without distinct striations, with semicircular Felisacus usingeri Woodward sclerite and distinct sclerotized rings laterally; Figures 7, 20 lateral oviducts placed almost medially, very Felisacus usingeri Woodward, 1954: 47 (original close to each other, spermathecal gland placed description). between lateral oviducts; dorsal labiate plate with distinct tubercles, without membranous lobe Diagnosis: Recognized by the following com- medially. bination of characters: head and pronotum mostly Distribution: Vanuatu (Tanna Is., Maewo yellow; antennal segment I yellow, brown apically; Is.) (fig. 20). transverse depression delimiting occipital region Host plants: Unknown. present only dorsally; labium reaching middle of Etymology: The species is named after mesosternum; distance between eye and prono- Tanna Is., where most of the examined speci- tum slightly longer than eye diameter. mens were collected. Description: Male. Unknown. Female. Total Discussion: Felisacus tanna is similar to F. length 3.7. COLORATION (fig. 7): Head: Mostly glabratus in coloration (figs. 5, 7) and the broad yellow. Labium: Uniformly yellow. Antenna: Seg- 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 151 ment I yellow, brown apically, segment II uni- Distribution: Philippines (Luzon) (fig. 20). formly brown. Eye brown. Thorax: Pronotum and Host plants: Unknown. scutellum yellow; thoracic pleura yellow; scent Discussion: Woodward (1954) described F. gland evaporative area whitish yellow. Hemelytron: usingeri from a single female, which we exam- Mostly translucent, colorless; clavus yellow; mark- ined, but did not dissect its genitalia. Felisacus ing along inner margin of corium brown; embo- usingeri is similar to F. auritulus, F. bradi, and F. lium with yellow tinge and pale brown margins; ovalau in coloration, broad antennal segment I, cuneus with yellow tinge, its outer margin brown. and dorsally elongate labial segment II. Felisacus Legs: Coxae whitish yellow; femora whitish yellow, auritulus differs from F. usingeri by the distance yellow apically; tibae yellow. Abdomen: Whitish from the eye to the pronotum, which is slightly yellow. SURFACE AND VESTITURE: Head, pro- shorter than the eye diameter, and antennal seg- notum, scutellum, clavus, and costal vein anteri- ment I dark brown with a yellow base. Felisacus orly with scarce setae subequal to or shorter than bradi differs from F. usingeri by the antennal seg- antennal segment II diameter; antennal segments ment I brown, whitish yellow basally and the I and II with suberect setae shorter than antennal labium reaching abdominal segment VIII. Feli- segment II diameter; femora clothed with suber- sacus ovalau can be separated from F. usingeri by ect setae slightly longer than antennal segment II antennal segment I yellow with a reddish tinge, diameter. STRUCTURE AND MEASURE- with red apically, and the presence of a pair of MENTS: Body ca. 4.7× as long as pronotum brown markings on the posterior part of the pro- width. Head: Depression delimiting occipital notum; in addition, the labium reaches abdomi- region distinct only dorsally; distance between nal segment V. depression and pronotum almost as long as eye Material examined: Holotype: PHILIP- diameter; longitudinal sulcus on dorsal surface PINES: Laguna: Los Banos, 14.16667°N shorter than eye diameter; distance from eye to 121.18333°E, 17 Jul 1936, R. Usinger, 1♀ pronotum slightly longer than eye diameter, not (00400378) (CAS). swollen; vertex ca. 1.8× as wide as eye, flat. Labium (as in Namyatova et al., 2016: figs. 6D, 9C): Reach- Felisacus vitilevu, sp. nov. ing middle of mesosternum; segment I and II combined subequal to segment III; segments I ca. Figures 7, 9N, 13V, W, 15L, 16I, J, 21 1.5× as long as wide; segment II twice as long as Diagnosis: Recognized by the following wide, elongate, dorsally; segment III distinctly combination of characters: pronotum mostly shorter than ventral side of head; segment IV pale brown to dark brown with reddish yellow or twice as long as segment III. Antenna: Segment I pale brown stripe medially, or at least humeral slightly widened, 1.3× as long as head width, 0.8× angles brown to dark brown; marking along as long as pronotum width; segment II 2.0× as inner margin of corium male to brown, not long as head width, 1.1× as long as pronotum extending toward R+M anteriorly and posteri- width. Thorax: Anterior part of pronotum dis- orly; antennal segment I pale brown or reddish tinctly shorter than posterior part; collar distinct; with yellow base, transverse depression on head posterior part of pronotum slightly raised; poste- present only dorsally; marking along inner mar- rior margin of pronotum slightly concave; prono- gin of corium narrow and short (fig. 7); antennal tum ca. 1.1× as wide as long and 1.7× as wide as segment I ca. 1.3–1.6× as long as head width; head; mesoscutum slightly exposed. Hemelytron: antennal segment II in male ca. 1.9–2.1× and in Area along inner margin of corium almost flat; female ca. 1.9–2.0× as long as head width; labium inner margin of cuneus convex (as in Namyatova reaching posterior margin of metasternum or et al., 2016: fig. 13F), medial margin ca. 3× as long slightly surpassing it; apex of ventral wall of geni- as base. Female genitalia not examined. tal capsule curved dorsally (fig. 15L); right para- 152 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 mere sickle shaped (fig. 13V); left paramere wide, dish yellow, sometimes yellow basally, dorsal with outgrowth posteriorly (fig. 13W); sclerotiza- surfaces red, rarely entire abdomen red. SUR- tion of ductus seminis around secondary gono- FACE AND VESTITURE: Corium smooth, pore twice as long as wide; vesical with four with shallow punctures. Dorsum with setae serrate spicules (fig. 9N). subequal to or slightly longer than antennal Description: Male. Total length 3.4–4.3. segment II diameter; segment II and femora COLORATION (fig. 7):Head: Mostly yellow, clothed with suberect setae shorter than anten- reddish yellow, or pale brown, tubercles around nal segment II diameter; abdomen clothed with antennal fossa whitish yellow and longitudinal short erect setae. STRUCTURE AND MEA- sulcus on dorsal surface reddish, sometimes SUREMENTS: Body ca. 4.3–4.8× as long as also with reddish marking dorsally or anteriorly pronotum width. Head: Depression delimiting above and between antennal fossa; clypeus often occipital region present only dorsally; distance reddish apically or uniformly reddish, buccula between depression and pronotum distinctly sometimes also reddish. Eye dark brown with shorter than eye diameter; longitudinal sulcus reddish tinge. Labium: Yellow, reddish yellow, on dorsal surface of head longer than eye diam- or pale brown, segment III sometimes darker eter; vertex ca. 1.8–2.2× as wide as eye, upraised than other segments. Antenna: Segment I pale (as in Namyatova et al., 2016: fig. 6D). Labium brown or reddish with yellow base; segment II (as in Namyatova et al., 2016: figs. 6D, 9C): pale brown to brown, often with reddish tinge, Reaching posterior margin of metasternum or sometimes darker than segment I; segments slightly surpassing it; segments I and II strongly III–IV brown to dark brown. Thorax: Pronotum shortened, combined longer than half of seg- mostly yellow, reddish yellow, or pale brown ment III; segment I shorter than wide; segment with posterior part often brown to dark brown, II slightly longer than wide, elongate dorsally; with yellow, reddish yellow, or pale brown lon- segment III longer than ventral side of head; gitudinal stripe medially, sometimes posterior segment IV 1.5× as long as segment III. part mostly reddish yellow to pale brown with Antenna: Segment I slightly widened, ca. 1.3– brown to dark brown humeral angles; scutellum 1.4× as long as head width, ca. 0.9–1.0× as long and mesoscutum reddish yellow to pale brown, as pronotum width; segment II ca. 1.9–2.1× as often with reddish tinge; thoracic pleura red- long as head width, ca. 1.3–1.4× as long as pro- dish yellow to pale brown; scent gland evapora- notum width; segment III slightly longer than tive area whitish yellow to yellow basally and segment II and twice as long as segment IV. orange or reddish apically, sometimes uniformly Thorax: Anterior part of pronotum somewhat reddish yellow to pale brown. Hemelytron: shorter than posterior part; collar delimited; Mainly translucent and colorless, marking along posterior part slightly raised; posterior margin inner margin of corium yellow to dark brown; of pronotum concave; pronotum ca. 1.1–1.3× as embolium with reddish or pale brown apex and wide as long and ca. 1.4–1.6× as wide as head; margins; cuneus with yellowish tinge, with yel- mesoscutum exposed. Hemelytron: Area along low outer margin and colorless inner margin; inner margin of corium almost flat; inner mar- membrane grayish or pale brown medially, cell gin of cuneus convex (as in Namyatova et al., yellow to brown, Legs: Coxae whitish yellow to 2016: fig. 13E), outer margin of cuneus ca. 3× yellow; femora whitish yellow to yellow basally as long as base. Abdomen: Genital capsule and darker, yellow, reddish yellow, or pale rotated left at right angle relative to rest of brown apically; tibiae and tarsus yellow, reddish abdomen. Genitalia: Genital capsule (fig. 15L) yellow, or pale brown, sometimes tarsal seg- ca. 1.5× as long as wide; ventral wall twice as ments II–III of fore- and middle tarsi brown. long as dorsal wall, with posterior margin of Abdomen: Ventral and lateral sides mostly red- ventral wall distinctly tapering apically, smooth, 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 153 without outgrowth(s), its apex slightly shifted spermathecal gland placed between lateral ovi- left, curved dorsally; sides of genital capsule not ducts; dorsal labiate plate with distinct tubercles, modified; paramere sockets more or less acute; without membranous lobe medially. distance between paramere sockets subequal to Distribution: Fiji Is. (Viti Levu Is.) (fig. 21). half of genital capsule width at base. Right para- Host plants: Unknown. mere (fig. 13V) sickle shaped; apex with small Etymology: The species is name after Viti toothlike outgrowth tubercle anteriorly, medial Levu Island, where all known specimens were part narrower than basal part, with setae, outer collected. margin convex and inner margin concave, with- Discussion: Felisacus vitilevu is very similar out swelling; outer angle indistinct; inner angle in coloration to F. madagascariensis and F. gres- distinct, not bearing setae; basal part only sitti (fig. 5). Felisacus madagascariensis can be slightly shorter than rest of paramere. Left para- separated from F. vitilevu by the anterior part of mere (fig. 13W) widened; apical part flattened, the pronotum brown, the labium reaching the with rectangular outgrowth on posterior side posterior margin of the mesosternum, antennal and without outgrowth on dorsal surface; mid- segment I reddish brown and ca. 1.7× as long as dle part widened dorsally, without swelling or the head width. Felisacus gressitti differs from F. outgrowth(s); setae placed on outer margin vitilevu by the clypeus mostly dark brown and medially. Aedeagus conjunctiva sclerotized only antennal segment II in both males and females is basally, secondary gonopore placed at base of ca. 1.6× as long as head width. The Fijian species endosoma in repose; sclerotized part of ductus F. ovalau is also very similar, but differs from F. seminis around secondary gonopore elongate, vitilevu in the pronotum mostly yellow, the but distinctly not reaching phallotheca mouth; labium reaching abdominal sternum VIII, the its dorsal wall longer than ventral one; vesica left paramere having three rounded outgrowths with four serrate spicules. (fig. 13F), and the sclerotization of the ductus Female. Total length 4.0–4.3. COLORATION seminis around the secondary gonopore as long (fig. 7): Head and labium: Similar to male, rarely as wide; the shape of the vesical sclerotization brown. Antenna: As in male. Thorax: Similar to also differs (cf. fig. 9G with fig. 9N). male, rarely posterior part of pronotum uniformly Material examined: Holotype: FIJI: Viti reddish yellow to pale brown. Legs and hemely- Levu: Korotongo, 18.17575°S 177.55112°E, 50 m, tron: As in male. Abdomen: Mostly as in male, Mar 1981, N.L.H. Krauss, 1♂ (00043142) sometimes uniformly pale brown. SURFACE (BPBM). Paratype: [[SHOULD THIS BE PLU- AND VESTITURE: As in male. STRUCTURE RAL?]]FIJI: Central Division: Bilo nr Lami, Viti AND MEASUREMENTS: Structure as in male; Levu Is., 18.11092°S 178.41011°E, 20 m, 27 Sep body ca. 4.2–4.4× as long as pronotum width; ver- 1979, M.K. Kamath, S.N. Lal, G.A and S.L. Sam- tex ca. 1.8–2.2× as wide as eye; antennal segment uelson, 1♀ (00043155) (BPBM). Additional I ca. 1.3–1.6× as long as head width, ca. 0.9–1.0× material: Lami, Viti Levu Is., 18.11092°S as long as pronotum width; segment II ca. 1.9– 178.41011°E, Feb 1951, N.L.H. Krauss, 1♀ 2.0× as long as head width, ca. 1.1–1.3× as long as (00043145), 3♂ (00043133–00043135) (BPBM); pronotum width; pronotum ca. 1.2× as wide as May 1951, N.L.H. Krauss, 2♂ (00043137, long and ca. 1.6–1.7× as wide as head. Genitalia 00043136) (BPBM); Nov 1957, N.L.H. Krauss, (fig. 16I, J): Dorsal labiate plate wider than dis- 3♂ (00043129–00043131), 1♀ (00043143) tance between apodemes of second valvulae, (BPBM). Lami, Viti Levu Is., 18.11092°S mostly smooth, without distinct striations; with 178.41011°E, 100 m, Dec 1978, N.L.H. Krauss, semicircular sclerite, widened posteriorly and dis- 1♂ (00043132), 1♀ (00043144) (BPBM). Mt tinct sclerotized rings laterally; lateral oviducts Koro’o, 1 km N South Ridge summit, 17.35°S placed almost medially, very close to each other, 177.55°E, 1000 m, 16 Nov 2005, C. Reid, 2♂ 154 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
(00018965, 00018966) (AM). Nukurua Forest, or uniformly brown. Eye brown with reddish Viti Levu Is., 17.91723°S 178.57739°E, 95 m, 15 tinge. Labium: Segments I–III yellow, segment IV Oct 1979, M.K. Kamath, G.A and S.L. Samuel- pale brown. Antenna: Segments I–II pale brown, son, 2♀ (00043156, 00042268t) (BPBM). [Mt segments III–IV brown to dark brown. Thorax: Kobalevu summit area], 432 m, 16 Oct 1979, Pronotum yellow with pale brown anterior mar- G.A. Samuelson, 1♀ (00045808) (BPBM). Viti gin; humeral angles and medial longitudinal stripe Levu: Korotongo, 18.17575°S 177.55112°E, 50 m, pale brown or brown, sometimes only posterior Mar 1981, N.L.H. Krauss, 3♀ (00043151– part of that stripe distinct; mesoscutum and scu- 00043153) (BPBM). Rdg. W of Nandarivatu, tellum yellow, punctures between them pale 17.61265°S 177.94588°E, 853 m, 11 Sep 1938, brown; thoracic pleura yellow to pale brown with E.C. Zimmerman, 1♀ (00043148) (BPBM). scent gland evaporative area yellow. Hemelytron: Western Division: Nadarivatu, Viti Levu Is., Mostly translucent; inner part of clavus opaque, 17.56°S 177.966°E, 850 m, 08 Mar 1963–13 Mar yellow to brown; outer part of clavus whitish yel- 1963, C.M. Yoshimoto, 1♂ (00043138), 2♀ low; corium whitish yellow, with narrow pale (00043149, 00043150) (BPBM). Nadarivatu, Viti brown or brown marking along inner margin of Levu Is., 17.56°S 177.966°E, Jan 1955, N.L.H. corium and base of cuneus, reaching R+M vein Krauss, 2♀ (00043146, 00043154), 3♂ posteriorly; embolium whitish yellow, brown api- (00043139–00043141) (BPBM). Nandarivatu, cally; cuneus whitish yellow; membrane pale Viti Levu Is., 17.56°S 177.96°E, 823 m, 10 Nov brown anteriorly and whitish apically; membrane 1964, N. McFarland, 1♂ (00017831) (SAMA). cell brown to dark brown. Legs: Coxae yellow; femora yellow, often reddish apically; middle and hind femora with reddish brown or brown mark- Felisacus wangae, sp. nov. ing in apical half; tibiae yellow with brown or dark Figures 7, 10M, 13X, Y, 15M, 16N, O, 21 brown longitudinal stripe at least basally; tarsal Diagnosis: Recognized by the following segment I yellow to pale brown, tarsal segments combination of characters: pronotum yellow II–III brown. Abdomen: Pale brown, brown dor- with three brown markings posteriorly; hemely- sally. SURFACE AND VESTITURE: Corium tron with narrow brown marking along inner smooth with scarce punctures. Dorsum with margin of corium, not extending toward R+M suberect setae longer than antennal segment II anteriorly (fig. 7); antennal segment I slightly diameter; antennal segment I with scarce suberect widened; occipital region delimited only dor- setae subequal to or longer than antennal segment sally; vertex upraised; dorsal surface of labial II diameter; femora with suberect setae longer segment I not elongate posteriorly (as in Namy- than antennal segment II diameter; abdomen atova et al., 2016: fig. 6E); right side of ventral clothed with short suberect setae. STRUCTURE wall of genital capsule widened (fig. 15M); right AND MEASUREMENTS: Body ca. 3.8–4.0× as paramere with distinct outer and inner angles long as pronotum width. Head: Depression delim- (fig. 13X); left paramere sickle shaped (fig. 13Y); iting occipital region present only dorsally; dis- sclerotization of ductus seminis around second- tance between this depression and pronotum ary gonopore twice as long as wide; vesica with shorter than eye diameter; longitudinal sulcus on irregularly shaped sclerite, bearing toothlike dorsal surface shorter than eye diameter; distance outgrowth and with small triangular spicule from eye to pronotum slightly longer than eye apically (fig. 10M). diameter, not swollen laterally; vertex ca. 1.4–1.5× Description: Male. Total length 4.6–4.7. as wide as eye, upraised. Labium: Reaching middle COLORATION (fig. 7): Head: Yellow to pale of mesosternum; segments I and II distinctly lon- brown, sometimes with dorsal surface brown; ger than wide, combined subequal to segment III; buccula brown anteriorly; clypeus brown apically segment II not elongate dorsally (as in Namyatova 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 155 et al., 2016: fig. 6E); segment IV as long as seg- teriorly, posterior angles brown; thoracic pleura ment III. Antenna: Segment I slightly widened, ca. pale brown with scent gland evaporative area yel- 1.2–1.4× as long as head width, ca. 0.8–1.0× as low basally and brown apically. Hemelytron: Sim- long as pronotum width; segment II ca. 1.6–1.7× ilar to male, clavus brown with brown margins, as long as head width, ca. 1.1× as long as prono- stripes on hemelytron brown. Legs: Similar to tum width; segment III slightly longer than seg- male, tibiae yellow with brown longitudinal ments II and IV. Thorax: Anterior part of stripe. Abdomen: Yellow to pale brown with pronotum shorter than posterior part; collar brown areas. SURFACE AND VESTITURE: As delimited; posterior part of pronotum slightly in male. STRUCTURE AND MEASUREMENTS: upraised; posterior margin of pronotum slightly Structure as in male. Genitalia (figs. 16N, O): concave; pronotum ca. 1.3× as wide as long and Dorsal labiate plate wider than apodemes of sec- ca. 1.6–1.7× as wide as head; mesoscutum slightly ond valvulae, with distinct striations, without exposed. Hemelytron: Area along inner margin of membranous ridge medially; semicircular scler- corium swollen; inner margin of cuneus convex ite and sclerotized rings absent; lateral oviducts (as in Namyatova et al., 2016: fig. 13E), outer mar- placed in posterior part, at distance from each gin of cuneus ca. 3× as long as base. Abdomen: other; spermathecal gland attached in posterior Genital capsule rotated at very small angle relative part, anteriorly to lateral oviducts; dorsal labiate to rest of abdomen. Genitalia: Genital capsule (fig. plate with membranous semicircular lobe anteri- 15M) twice as long as wide; ventral wall ca. 1.5× orly, with tubercles only along posterior margin as long as dorsal wall, its posterior margin smooth, of posterior wall. semioval, without outgrowth, its apex placed Distribution: Borneo (Sarawak, Kaliman- medially; right side of genital capsule curved, left tan) (fig. 21). side not widened; paramere sockets more or less Host plants: Unknown. acute; distance between paramere sockets sub- Etymology: The species is name after Xiao- equal to half of genital capsule width at base. jing Wang from Guo-Qing Liu’s laboratory Right paramere (fig. 13X) sickle shaped; apex con- (Nankai University) who spent the better part of cave posteriorly; medial part as wide as basal part, a year at the University of New South Wales and without setae, with outer margin convex and helped us by translating species descriptions inner margin with slightly widened; outer angle from Chinese. distinct, inner angle distinct, without outgrowth(s) Discussion: The sole female specimen has or swelling, bearing setae; basal part as long as rest glue on the dorsum, and segments II–IV are of paramere. Left paramere (fig. 13Y) sickle missing. Felisacus wangae is not similar to any shaped; apical part not flattened, without tooth or species of the genus. It is the sister species of F. outgrowth on apical part; middle side slightly curvatus, as they both possess similar female widened, without swelling or outgrowth(s); setae genitalia with distinct striations (cf. fig. 16L, N), placed on middle part near outer margin. a broad right side of the genital capsule (cf. figs. Aedeagus conjunctiva not sclerotized, secondary 14J, 15M), and an apical sclerotized part of the gonopore placed at base of vesical in repose; scler- ductus seminis twice as long as wide (cf. fig. 10E, ite surrounding secondary gonopore twice as long M). Felisacus curvatus differs from F. wangae in as wide; vesica with two spicules; one of them very the marking on the hemelytron broad (fig. 4), the small, smooth; second one large, bearing toothlike labium only slightly surpassing the posterior outgrowths (fig. 10M). margin of the prosternum, and the presence of Female. COLORATION (fig. 7): Head: Pale two serrate spicules, bearing toothlike out- brown with clypeus and marking on buccula growths (fig. 10E). brown. Thorax: Similar to male, but medial stripe Material examined: Holotype: INDONE- on posterior part of pronotum distinct only pos- SIA: Kalimantan Timur: Pasir Dist.: 4 km W 156 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
Lumut, Gunung Lumut, 1.46305°S 115.99815°E, with grayish tinge; membrane cells yellow with 380 m, 15 Nov 2005, E. Gasso Miracle, light trap, reddish tinge. Abdomen: Yellow with reddish 1♂ (00017870) (NML). Paratypes: INDONE- tinge. SURFACE AND VESTITURE: Corium SIA: Kalimantan Timur: Pasir Dist.: 4 km W smooth, without punctures. STRUCTURE AND Lumut, Gunung Lumut, 1.46305°S 115.99815°E, MEASUREMENTS: Body ca. 4.2–4.3× as long as 380 m, 15 Nov 2005, E. Gasso Miracle, Light pronotum width. Head: Depression delimiting Trap, 1♀ (00017869) (NML). MALAYSIA: Sar- occipital region present dorsally and laterally; awak: Nanga Pelagus, 2.01666°N 112.93305°E, distance between depression and pronotum dis- 07 Aug 1958–14 Aug 1958, T.C. Maa, 1♂ tinctly shorter than eye diameter; longitudinal (00043874) (BPBM). Nanga Pelagus nr Kapit, sulcus on dorsal surface longer than eye diame- 2.01666°N 112.93305°E, 585 m, 07 Aug 1958–14 ter; distance from posterior margin of eye to pro- Aug 1958, T.C. Maa, 1♂ (00043875) (BPBM). notum as long as eye, not swollen laterally; vertex ca. 2.0× as wide as eye, upraised (as in Namya- tova et al., 2016: fig. 6D). Labium (as in Namya- Felisacus webbi, sp. nov. tova et al., 2016: figs. 6D, 9C ): Reaching Figures 7, 10N, 13Z, AA, 15N, 21 abdominal segment V; segments I and II com- Diagnosis: Recognized by the following bined subequal to half of segment III; segment I combination of characters: head and pronotum shorter than wide; segment II slightly longer yellow, pronotum with pair of brown markings than wide, its dorsal surface elongate posteriorly; near humeral angles (fig. 7); transverse depres- segment IV twice as long as segment III. Antenna: sion delimiting occipital region present only dor- Segment I cylindrical (as in Namyatova et al., sally; antennal segment I slightly widened; 2016: fig. 8A), ca. 1.5–1.6× as long as head width, labium reaching abdominal segment V; dorsal ca. 0.9–1.0× as long as pronotum width; segment surface of labial segment I elongate posteriorly II ca. 1.9–2.1× as long as head width, ca. 1.2– (as in Namyatova et al., 2016: figs. 6D, 9C); ven- 1.3× as long as pronotum width. Thorax: Ante- tral wall of genital capsule with outgrowth from rior part of pronotum shorter than posterior right-hand side dorsally, its apex not curved dor- part; collar delimited; posterior part of pronotum sally (fig. 15N); both parameres narrow, sickle slightly upraised; posterior margin of pronotum shaped, without outgrowths; right paramere concave, pronotum ca. 0.8× as wide as long and swollen medially; outer angle of left paramere ca. 1.6× as wide as head; mesoscutum exposed. present, inner angle rounded (fig. 13Z, AA); Hemelytron: Area along inner margin of corium vesica with long curved spicule (fig. 10N). almost flat; inner margin of cuneus convex (as in Description: Male. Total length 3.6–3.7. Namyatova et al., 2016: fig. 13E), outer margin of COLORATION (fig. 7): Head: Yellow, frons and cuneus ca. 3× as long as base. Abdomen: Genital clavus with reddish tinge. Labium: Uniformly capsule rotated at right angle relative to other yellow. Antenna: Segment I yellow with reddish segments. Genitalia: Genital capsule (fig. 15N) stripe and apex, segments II–IV reddish brown. twice as long as wide; ventral wall ca. 1.3× as Thorax: Pronotum yellow with pair of dark long dorsal wall, with posterior margin of ventral brown markings posteriorly; scutellum uni- wall rounded, not folded; right side folded, left formly yellow; thoracic pleura and scent gland side not widened; paramere sockets more or less evaporative area uniformly yellow. Hemelytron: acute; distance between paramere sockets ca. Mostly colorless, transparent; clavus whitish yel- 0.3× as long as genital capsule base. Right para- low to brown; corium with reddish tinge poste- mere (fig. 13Z) distinctly curved in apical half; riorly, marking along inner margin of corium anterior part straight posteriorly; medial part yellow; embolium with yellow tinge, reddish api- slightly wider than basal part, its outer margin cally; cuneus yellow along margins; membrane concave and inner margin widened; outer angle 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 157 distinct; inner angle rounded; basal part sub- on corium distinctly C-shaped, reaching R+M equal to half of rest of paramere. Left paramere anteriorly and posteriorly, anterior part of this (fig. 13AA) sickle shaped, apical part without marking inclined posteriorly (fig. 7); antennal outgrowth(s); middle part slightly widened, segment II widened (as in Namyatova et al., bearing setae, its inner margin without swelling 2016: fig. 8B), red; vertex flat dorsally (as in or outgrowth. Aedeagus (fig. 10N) conjunctiva Namyatova and Cassis, in press: fig. 6E) and membranous; with single large curved spicule; straight laterally; distance between transverse secondary gonopore placed at base of vesical in depression on head and pronotum longer than repose; sclerotization of ductus seminis sur- eye diameter; labium reaching posterior margin rounding secondary gonopore shorter than wide; of metasternum; cuneus uniformly colorless; api- vesica with single large curved spicule. cal half of femora red; ventral wall of genital cap- Female. Unknown. sule with toothlike outgrowth posteriorly (fig. Distribution: Madagascar (fig. 21). 15O); right paramere sickle shaped, without out- Host plants: Unknown. growth on inner angle (fig. 13AB); apical part of Etymology: The species is names after Mick left paramere with toothlike outgrowth apically Webb, the curator in the Natural History and outgrowth on dorsal surface; and medial Museum who helped the first author (A.A.N.) part of left paramere with distinct outgrowth (fig. during her visit to the museum and loaned speci- 13AC); apical half of ductus seminis sclerotized, mens of the genus Felisacus, including two speci- its apex hooked (fig. 10O). mens of this new species. Description: Male. Total length 3.3–3.5. Discussion: Felisacus webbi is most similar to COLORATION (fig. 7): Head: Mostly yellow F. ovalau in coloration and structure (figs. 6, 7). with reddish markings. Eye brown with reddish The latter species differs from F. webbi in the tinge. Labium: Uniformly yellow. Antenna: Seg- ventral wall of the genital capsule curved api- ment I red brown apically and often with yellow cally, without an outgrowth on the right-hand base; segments II yellow to brown, sometimes side (fig. 15F), the left paramere having three yellow, brown apically; segments III–IV yellow. rounded outgrowths (fig. 13F), and the vesica Thorax: Anterior part of pronotum yellow to pale having a number of small toothlike spinules and brown with reddish tinge or red; posterior part two large spicules (fig. 9G). of pronotum uniformly dark brown; scutellum Material examined: Holotype: MADAGAS- and mesoscutum uniformly dark brown; tho- CAR: Fianarantsoa: Ranomafana National Park, racic pleura dark brown, metepimeron whitish Setam Lodge Hotel, 21.2501°S 47.42678°E, 917 yellow; scent gland evaporative area whitish yel- m, 04 Nov 2005–21 Nov 2005, G. Martin, D.L.J. low. Hemelytron: Clavus opaque, brown anteri- Quicke and L.P. Holland, 1♂ (00019536) orly, whitish posteriorly with brown or reddish (BMNH). Paratype: MADAGASCAR: Fianara- margins; corium, embolium, cuneus, and mem- ntsoa: Ranomafana National Park, Setam Lodge brane mostly transparent, colorless; corium with Hotel, 21.2501°S 47.42678°E, 917 m, 04 Nov brown to dark brown posterior angle and with 2005–21 Nov 2005, G. Martin, D.L.J. Quicke and C-shaped dark brown marking, reaching R+M L.P. Holland, 1♂ (00019537) (BMNH). anteriorly and posteriorly, anterior part of this marking inclined posteriorly; embolium brown apically; cuneus with yellow outer margin; mem- Felisacus yasunagai, sp. nov. brane cell colorless. Legs: Coxae whitish yellow to yellow; femora whitish yellow to yellow basally Figures 7, 10O, 13AB, AC, 15O, 21 and reddish apically, often with brown marking Diagnosis: Recognized by the following medially; tibiae brown basally and whitish yellow combination of characters: dark brown marking to yellow apically; tarsi whitish yellow to yellow. 158 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
Abdomen: Whitish yellow, sometimes with rest of abdomen. Genitalia: Genital capsule (fig. brown markings. SURFACE AND VESTITURE: 15O) ca. 1.5× as long as wide; ventral wall ca. Corium with distinct punctation. Dorsum 1.5× as long as dorsal wall, with posterior margin clothed with setae subequal to or shorter than of ventral not curved dorsally, with small tooth- antennal segment II diameter, those setae longer like outgrowth; right side of genital capsule on head and posterior part of pronotum; anten- folded, left side widened; paramere sockets dis- nal segment I clothed with suberect setae shorter tinctly acute; distance between paramere sockets than antennal segment II diameter; femora ca. 0.15–0.2× as long as genital capsule width at clothed with suberect setae longer than antennal base. Right paramere (fig. 13AB) sickle shaped; segment II diameter; abdomen clothed with apex slightly concave; medial part narrower than setae of different length. STRUCTURE AND basal part, without setae, outer margin of medial MEASUREMENTS: Body ca. 3.9–4.2 × as long part convex, inner margin concave, with swell- as pronotum width. Head: Transverse depression ing; outer angle absent, inner margin present, delimiting occipital region present only dorsally; swollen, without setae or outgrowth(s); basal distance between depression and pronotum lon- part subequal to rest of paramere. Left paramere ger than eye diameter; longitudinal sulcus on (fig. 13AC) apical part not flattened, with tooth- dorsal surface of head shorter than eye diameter; like outgrowth apically and with outgrowth on distance between eye and pronotum longer than dorsal surface; middle part with outgrowth at eye diameter, not swollen; vertex ca. 1.8–2.1× as inner side, without setae. Aedeagus (general view wide as eye, flat (as in Namyatova et al., 2016: fig. as Namyatova et al., 2016: fig. 22M) conjunctiva 6E). Labium: Reaching posterior margin of membranous; vesica without spicules; apical half metasternum or slightly surpassing it, segments of ductus seminis sclerotized, secondary gono- I and II shorter than width (as in Namyatova et pore placed near phallotheca mouth in repose; al., 2016: fig. 6E), combined subequal to half of ductus seminis hooked apically (Namyatova et segment III; dorsal surface of segment II not al., 2016: fig. 10O). elongate dorsally (as in Namyatova et al., 2016: Female. Total length 3.5–3.8. COLORATION, fig. 6E), segment III shorter than ventral side of SURFACE, AND VESTITURE (fig. 7): As in head; segment IV twice as long as segment III. male. STRUCTURE AND MEASUREMENTS: Antenna: Segment I distinctly longer than head Structure as in male; body ca. 3.8–4.1× as long as width, swollen (as in Namyatova et al., 2016: fig. pronotum width; vertex ca. 1.6–1.9× as wide as 8B), ca. 1.3–1.6× as long as head width, ca. 0.7– eye; antennal segment I ca. 1.2–1.4× as long as 0.8× as long as pronotum width; segment II ca. head width, ca. 0.6–0.7× as long as pronotum 1.4–1.5× as long as head width, ca. 0.8× as long width; segment II ca. 1.3–1.5× as long as head as pronotum width; segments III almost twice as width, ca. 0.6–0.8× as long as pronotum width; long as segment II and twice as long as segment pronotum ca. 1.1–1.2× as wide as long and ca. IV. Thorax: Anterior part of pronotum distinctly 1.8–2.2× as wide as head. Genitalia: Dorsal labi- shorter than posterior part; collar distinct; pos- ate plate small and transparent, as wide as dis- terior part slightly upraised; posterior margin of tance between apodemes of second valvula, pronotum straight or concave; pronotum ca. without striations or membranous ridge medi- 1.1–1.2× as wide as long and ca. 1.8–1.9× as wide ally; semicircular sclerite and sclerotized rings as head; mesoscutum usually exposed. Hemely- absent; lateral oviducts placed medially, close to tron: Area along inner margin of corium swollen; each other, far from lateral margins of dorsal inner margin of cuneus straight (as in Namya- labiate plate (fig. 16K); dorsal labiate plate with- tova et al., 2016: fig. 13F), outer margin of cuneus out distinct tubercles, with triangular membra- twice as long as base. Abdomen: Genital capsule nous lobe posteriorly (as in fig. 16H). rotated left at almost right angle relative to the Distribution: Philippines (Luzon Is.) (fig. 21). 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 159
Host plant: Unknown. eye diameter; antennal segment I red; antennal Etymology: Species is named after Tomo- segment II reddish brown; labium reaching pos- hide Yasunaga, who loaned us specimens of Feli- terior margin of mesosternum; cuneus translu- sacus from his collection. cent, red apically; dorsal wall of genital capsule Discussion: Place of attachment of sperma- with tooth posteriorly (fig. 15P); right paramere thecal gland could not be seen. Felisacus yasuna- sickle shaped with curved outgrowth on inner gai is similar to Felisacus species with a developed angle (fig. 13AD); apical part of left paramere C-shaped marking on corium, but those species narrow, with tooth apically and outgrowth dor- differ in the anterior part of the C-shaped mark- sally; medial part with wide outgrowth (13 AE); ing not inclined posteriorly; also the labium is half of ductus seminis sclerotized, hooked api- shorter, varying from reaching the posterior cally, vesica with small spicule (fig. 10P). margin of prosternum to the posterior margin of Description: Male. Total length 3.4–4.1. the mesosternum. COLORATION (fig. 7): Head: Mostly yellow to Material examined: Holotype: PHILIP- pale brown, with reddish tinge or reddish mark- PINES: Camarines Sur: Iriga Mt, 13.45416°N ings, dorsal and lateral sides often darker than 123.45055°E, 26 Apr 1962, H.M. Torrevillas, 1♂ frontal and ventral surfaces; clypeus sometimes (00043225) (BPBM). Paratypes: PHILIPPINES: uniformly reddish; buccula sometimes brown. Camarines Sur: Iriga Mt, 13.45416°N Eye brown to dark brown with reddish tinge. 123.45055°E, 26 Apr 1962, H.M. Torrevillas, 6♂ Labium: Segments I–II brown, segments III–IV (00043212–00043217), 5♀ (00043220– yellowish to pale brown, sometimes uniformly 00043224) (BPBM). Isarog Mt, 20 km E of Naga, yellow to pale brown. Antenna: Segment I red, 13.65805°N 123.37305°E, 550 m, 06 Apr 1963, sometimes brown apically, segment II reddish H.M. Torrevillas, Light Trap, 1♀ (00043341) brown, often apically darkened, segment III pale (BPBM). Isarog Mt, Pili, 13.65805°N 123.37305°E, brown to brown. Thorax: Anterior part of pro- 600 m, 05 Apr 1965, H.M. Torrevillas, 1♂ notum pale brown, often brown around fore- (00043219), 1 sex unknown (00043226) (BPBM). coxa, sometimes with reddish tinge, with brown Mt. Isarog, Pili, 13.70805°N 123.75083°E, 800 m, to dark brown anterior margin; posterior part of 27 Apr 1965, H.M. Torrevillas, 1♂ (00043218) pronotum dark brown; scutellum and mesoscu- (BPBM). Luzon: Salton Pass, Nueva Vizcaya, tum uniformly brown to dark brown; thoracic 15.11666°N 120.95°E, 915 m, 09 Apr 1968–10 pleura brown to dark brown; scent gland evapo- Apr 1968, D.E. Hardy, 1♀ (00043342) (BPBM). rative area whitish yellow to pale brown, some- Misamis Oriental: Mt. Kibungol, 09 Apr 1960– times brown apically. Hemelytron: Mostly 18 Apr 1960, H.M. Torrevillas, light trap, 1♀ translucent; clavus opaque, uniformly dark (00045817) (BPBM). brown; corium translucent, colorless with brown anterior angles and C-shaped dark brown mark- ing reaching R+M anteriorly and posteriorly, its Felisacus zuparkoi, sp. nov. anterior not inclined posteriorly; embolium Figures 7, 10P, 13AD, AE, 15P, 21 with brown outer margin and colorless inner Diagnosis: Recognized by the following margin, brown apex and sometimes pale brown combination of characters: presence of dark or reddish marking medially; cuneus colorless brown C-shaped marking on corium, reaching with reddish apex and pale brown margins; R+M anteriorly and posteriorly, its anterior part membrane gradually changing color from pale not inclined posteriorly (fig. 7); vertex flat (as in brown to brown anteriorly and pale brown or Namyatova et al., 2016: fig. 6E), not widened lat- whitish posteriorly, cell pale brown to brown. erally behind eye; distance between transverse Legs: Coxae whitish yellow to yellow; femora depression on head and pronotum longer than whitish yellow to yellow often with pale brown 160 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 or reddish apices; tibiae yellow, often with pale tova et al., 2016: fig. 13F), outer margin of brown or red bases, rarely also with pale brown cuneus twice as long as base. Abdomen: Genital apices; tarsi uniformly whitish yellow to yellow. capsule rotated at small angle relative to rest of Abdomen: Yellowish with pale brown to brown abdomen. Genitalia: Genital capsule (fig. 15P) dorsal surface and segments VIII–IX, some- ca. 1.5× as long as wide; ventral wall ca. 1.5× as times reddish apically, abdomen sometimes uni- long as dorsal wall, with posterior margin of formly pale brown. SURFACE AND ventral wall not curved dorsally, with toothlike VESTITURE: Corium smooth, with scarce and outgrowth; right side of genital capsule folded, shallow punctation. Dorsum and femora with left side not widened; paramere sockets dis- setae subequal to or longer than antennal seg- tinctly acute; distance between paramere sockets ment II diameter; antennal segment I clothed ca. 0.25× as long as genital capsule width at with suberect setae shorter than antennal seg- base. Right paramere (fig. 13AD) sickle shaped; ment II diameter; abdomen clothed with short apex straight posteriorly; medial part narrower erect setae. STRUCTURE AND MEASURE- than basal part, without setae, outer margin of MENTS: Body ca. 3.9–4.2× as long as pronotum medial part convex and inner margin concave; width. Head: Depression delimiting occipital outer angle absent; inner angle present, with region present only dorsally; distance between curved outgrowth bearing setae; basal part depression and pronotum as long as eye diam- slightly shorter than rest of paramere. Left para- eter; longitudinal sulcus on dorsal surface of mere (fig. 13AE) distinctly curved medially; api- head shorter than eye diameter; distance from cal part not flattened, with toothlike outgrowth head to pronotum longer than eye diameter, not on posterior side apically, without other out- swollen; vertex ca. 1.6–1.7× as wide as eye, flat growths; inner margin of middle part with (as in Namyatova et al., 2016: fig. 6E). Labium: broad outgrowth with a number of toothlike Reaching posterior margin of mesosternum or outgrowths, bearing setae. Aedeagus (general slightly surpassing it; segment I and II combined view as in Namyatova et al., in press: fig. 22M) subequal to half of segment III; segment I as conjunctiva membranous; vesica with single long as wide; segment II slightly longer than small claw-shaped spicule; apical half of ductus width not elongate dorsally (as in Namyatova et seminis sclerotized, secondary gonopore placed al., 2016: fig. 6E); segment III as long as ventral near phallotheca mouth in repose; ductus semi- side of head; segment IV ca. 1.5× as long as seg- nis hooked apically (fig. 10P). ment III. Antenna: Segment I slightly longer Female. Total length 3.3–3.4. COLORATION than head width, swollen (as in Namyatova et (fig. 7): Similar to male, cuneus rarely whitish al., 2016: fig. 8B), ca. 1.1–1.2× as long as head with pale brown outer margin. SURFACE AND width, ca. 0.6–0.7× as long as pronotum width; VESTITURE: As in male. STRUCTURE AND segment II ca. 1.6–1.7× as long as head width, MEASUREMENTS: Structure as in male; body ca. 0.9–1.0× as long as pronotum width; seg- ca. 3.7–3.9× as long as pronotum width; vertex ment III slightly longer than segment II. Thorax: ca. 2.0–2.2× as wide as eye; antennal segment I Anterior part of pronotum shorter than poste- ca. 1.1× as long as head width, ca. 0.6× as long as rior part; collar delimited; posterior part of pro- pronotum width; segment II ca. 1.5× as long as notum slightly upraised; posterior margin of head width, ca. 0.9× as long as pronotum width; pronotum straight or slightly concave; prono- pronotum ca. 1.2–1.4× as wide as long and ca. tum ca. 1.1–1.3× as wide as long and ca. 1.6– 1.8–2.0× as wide as head. Genitalia: Dorsal labi- 1.8× as wide as head; mesoscutum slightly ate plate very small and transparent, as wide as exposed, sometimes not exposed; Hemelytron: distance between apodemes of second valvulae, Area along inner margin of corium swollen; without striations or membranous ridge medi- inner margin of cuneus straight (as in Namya- ally; semicircular sclerite and sclerotized rings 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 161 absent; lateral oviducts placed in posterior part; 1958, T.C. Maa, 1♂ (00043183) (BPBM). Pang- spermathecal gland attached near anterior mar- kalan Tebang, 350 m, 05 Sep 1958, T.C. Maa, 3♀ gin (as in fig. 16G); dorsal labiate plate without (00043189, 00043190, 00043343) (BPBM). San- distinct tubercles, without membranous lobe tubong, Kuching, 1.71666°N 119.3°E, 1150 m, 18 posteriorly (as in fig. 16B). Jun 1958–30 Jun 1958, T.C. Maa, 3♂ (00043180, Distribution: Borneo (Sarawak), Philip- 00043181), 1♀ (00043181) (BPBM). PHILIP- pines (Mindanao Is.) (fig. 21). PINES: Mindanao: [Mt. Empagatao], 1125 m, Host plants: Unknown. 19 Apr 1961–30 Apr 1961, H.M. Torrevillas, 1♂ Etymology: The species is named after (00045809) (BPBM). Robert Zuparko, curatorial assistant at the Cali- fornia Academy of Sciences, for loaning mate- ACKNOWLEDGMENTS rial of Felisacus. Discussion: Antennal segment IV is lost in This work was part of a Ph.D. project by one both males and females. Felisacus zuparkoi is of us (A.A.N.), and was supported by a UNSW similar to F. longiceps and F. magnificus exter- International Scholarship and a UNSW Ecology nally. Both those species differ from F. zuparkoi and Evolutionary Research Centre Postgraduate in the absence of the vesical spicules (fig. 10H, J) Writing and Skills Transfer Award. The project and the narrow outgrowth on the left paramere was also sponsored by RFBR research grant no. (fig. 12R, X). Felisacus longiceps also can be sepa- 15-29-02533 ofi_m. We are also grateful to the rated by the antennal segment II often yellow, Royal Zoological Society (London), Ernst Mayr and F. magnificus differs in the shape of the left Travel Grant in Animal Systematics (Harvard paramere, with the apical part flattened (fig. University, Museum of Comparative Zoology), 12X). International Heteropterists’ Society and UNSW Felisacus zuparkoi is also similar to Felisacus School of Biological, Earth and Environmental ceylonicus and F. lindbergae in coloration (figs. 4, Sciences who provided travel funds for one of us 5). F. ceylonicus differs from F. zuparkoi in having (A.A.N.) to examine specimens preserved in a swollen vertex behind the eyes and the narrow non-Australian museums. The Australian Bio- outgrowth on the left paramere (fig. 11S). Feli- logical Resources Study (ABRS) provided a grant sacus lindbergae can be separated by the labium to present results on Bryocorinae classification in reaching the posterior margin of the prosternum New Zealand, at the 3rd Combined Australian or slightly surpassing it and the outgrowth on the and New Zealand Entomological Societies Con- left paramere is short (fig. 12N). ference. Much of this material was assembled Material examined: Holotype: MALAYSIA: and originally examined by one of us (G.C.) Sarawak: Bau District: Bidi, Bau District, under an ABRS grant to him. 1.38389°N 110.13357°E, 165 m, 03 Sep 1958, T.C. We thank the curators of the following muse- Maa, 1♂ (00043182) (BPBM). Paratypes: ums for loaning the material: Dave Britton (AM); MALAYSIA: Sabah: Kalabakan, 4.4167°N Randall Schuh (AMNH); Tom Weir and Beth 117.4833°E, 08 Nov 1958–15 Nov 1958, T.C. Mantle (ANIC); Shepherd Myers (BPBM); Robert Maa, 1♀ (00043191) (BPBM); 10 Nov 1958–19 Zuparko (CAS); David Redei and András Orosz Nov 1958, T.C. Maa, 1♂ (00043188) (BPBM). (HNHM); Eliane De Coninck and Marc de Meyer Ranau, 5.9667°N 116.6833°E, 28 Sep 1958–30 (ISNB); Larry Huldén (MZH); Ken Walker Sep 1958, T.C. Maa, 1♂ (00043187) (BPBM). (MVMA); Gunvi Lindberg (NHRS); Guoqing Liu Sarawak: Bau District.: Bidi, Bau District, (NKMU); Mei-Ling Chan (NMNS); Michael 1.38389°N 110.13357°E, 03 Sep 1958, T.C. Maa, Braby (NTM); Christine Lambkin and Susan 2♂ (00043184, 00043185) (BPBM). Bidi, Bau Wright (QM); Peter Hudson (SAMA); Katrina District, 1.38389°N 110.13357°E, 165 m, 03 Sep Menard and Ed Riley (TAMU); Tomohide Yasu- 162 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403 naga (Japan); Brad Balukjian (UCB); Thomas Catalogue of Australia, vol. 27.3A. Melbourne, Aus- Henry, Jr. (USNM); Terry Houston (WAMP), tralia: CSIRO. Dmitry Gapon (ZISP), Ian Millar (PPRI) and vari- Cassis, G., and R.T. Schuh. 2012. Systematics, biodiver- ous curators from NML. We also thank Mick sity, biogeography, and host associations of the Miridae (Insecta: Hemiptera: Heteroptera: Miridae). Webb, Laurence Livermore, and Vladimir Bla- Annual Review of Entomology 57: 377–404. goderov (BMNH) for assistance with working in China, W.E. 1944. New and little known West African the collection and imaging of the specimens in the Miridae (Capsidae) (Hemiptera-Heteroptera). Bul- Sackler Biodiversity Imaging Lab. We thank Fedor letin of Entomological Research 35: 171–191. Konstantinov (St. Petersburg State University) for Christenhusz, M.J., and Chase M.W. 2014. Trends and help with access to specimens and use of facilities. concepts in fern classification. Annals of Botany We thank all the people from the Cassis’ lab 113: 571–594. (UNSW), especially Celia Symonds for technical Davis, N.T. 1955. 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APPENDIX 1 7 Labial segment II length: 0, shorter than width, as long Characters and Character States Used in as wide, or only slightly longer than wide (Namyatova et al., in press: fig. 6D, E); 1, elongate, distinctly longer the Phylogenetic Analysis than wide (Namyatova et al., in press: fig. 9A, E). In most species of Felisacus state 0 is present, except for Head F. curvatus and F. wangae, which possess the state 1. 0 Dorsal side of head posteriorly: 0, flat (Namyatova et In outgroup only state 1 is present. al., in press: fig. 6E); 1, at least slightly swollen 8 Dorsal side of labial segment II shape: 0, not elongate (Namyatova et al., in press: fig. 6D). Most of the dorsally (Namyatova et al., in press: fig. 6E); 1, Felisacus species have swollen group; in outgroup elongate dorsally (Namyatova et al., in press: fig. both states occur. 6D). In Felisacus species both states are present; in 1 Longitudinal depression on dorsal side of head: 0, outgroup species only state 0 occurs. absent or very shallow; 1, distinctly shorter than 9 Labial segment III relative length: 0, distinctly eye diameter; 2, as long as or longer than eye shorter than ventral side of head; 1, subequal or diameter (Namyatova et al., in press: fig. 4E). In longer to ventral side of head (Namyatova et al., in Felisacus states 1 and 2 are present; in outgroup press: fig. 9C). In Felisacus and outgroup species species states 0 and 1 occur. both states occur. 2 Transversal depression, delimiting occipital region: 0, 10 Labial segment IV relative length: 0, as long as absent or very shallow; 1, distinct only dorsally segment III; 1, distinctly longer than segment III. between eyes; 2, distinct, surrounding neck. In State 1 is present in Felisacus and Pachypeltis reuteri. Felisacus states 1 and 2 are present; in outgroup states 0 and 2 occur. Thorax 3 Distance between transversal depression and 11 Calli position: 0, separate; 1, fused (Namyatova et pronotum: 0, distinctly shorter than eye diameter; al., in press: fig. 4A, E). Calli fused in all species of 1, as long as or longer than eye diameter. This Felisacus. Within outgroup species they are fused character was not coded for Stenotus binotatus and only in Pachypeltis reuteri. Nesidiocoris tenuis, where transversal depression is 12 Callosite region delimitation: 0, not delimited; 1, absent. State 0 is present in most of the Felisacus delimited (Namyatova et al., in press: fig. 4A, E). species as well as in Pachypeltis reuteri. State 1 is present in all species of Felisacus and 4 Bucculae shape posteriorly: 0, not merged posteriorly Pachypeltus reuteri. (Namyatova et al., in press: fig. 6A, G); 1, merged 13 Punctures on sulcus between mesoscutum and scutellum posteriorly (Namyatova et al., in press: fig. 6D, E). occurrence: 0, absent; 1, present. State 1 is present in Buccula are merged posteriorly in all species of all species of Felisacus and Pachypeltus reuteri. Felisacus. It is not merged in species included in the 14 Posterior part of pronotum coloration: 0, yellow to outgroup in this analysis, however, within subfamily pale brown, sometimes dark brown with yellow to Bryocorinae it is also merged in the tribe Bryocorini pale brown stripe; 1, mostly yellow to pale brown, (Namyatova et al., in press: fig. 6F). with stripe along posterior margin; 2, uniformly 5 Antennal segment I shape: 0, straight (Namyatova et brown to black. In Felisacus all states are present, al., in press: fig. 8A); 1, widened basally or medially but state 0 is most common. In ougroups species (Namyatova et al., in press: fig. 8B). In Felisacus, as only state 0 is present. well as in outgroup species, both states occur. 15 Suture between meso- and metapleuron: 0, inferiorly 6 Labial segment I length: 0, shorter or as long as obsolete, mesepimeron and metepisternum fused width (Namyatova et al., in press: fig. 6D, E); 1, ventrad to metathoracic spiracle (Namyatova et al., elongate, twice as long as wide; 2, elongate, more in press: fig. 14A, B); 1, complete (Namyatova et al., than twice as long as wide (Namyatova et al., in in press: fig. 14C). The state 0 is present in Felisacus press: fig. 9A, E). In most species ofFelisacus state and Pachypeltis reuteri. In outgroup species state 1 0 is present, except for F. curvatus, F. philippinensis, is present. Within Bryocorinae in all Monaloniini and F. wangae which possess the state 1. In outgroup the suture between meso- and metapleuron inferi- only state 2 is present. orly obsolete (see Namyatova et al., in press). 166 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
16 Setae on hemelytron density: 0, present only 26 Small tooth on posterior margin of genital capsule anteriorly; 1, present, covering entire hemelytron. occurrence: 0, absent; 1, present (e.g., fig. 14T, V, Y). The state 0 is present in all Felisacus species, all State 1 is present in some species of Felisacus. outgroup species state 1 occurs. 27 Apex of genital capsule direction: 0, directed poste- 17 Row of punctures on clavus occurrence: 0, absent riorly, not curved; 1, directed dorsally, curved (figs. (Namyatova et al., in press: fig. 12C); 1, present 14X, 15F, L). State 1 is present in F. luzonus, F. ova- (Namyatova et al., in press: fig. 11A, C). State 1 is lau, and F. vitilevu. present in Felisacus and Pachypeltis reuteri. 28 Inner margin of left paramere socket shape: 0, 18 Inner part of clavus colouration: 0, whitish yellow to rounded or slightly angulate (e.g., figs. 14A, L, 15K, pale brown, usually transparent; 1, brown to black, L0; 1, distinctly angulate (e.g., fig. 14T, V, X). In usually matt. In Felisacus and outgroup species both Felisacus both states are present. In outgroup spe- states are present. cies only state 0 is present. 19 Corial fracture length: 0, short, almost indistinct 29 Relative size of parameres: 0, right paramere lon- (Namyatova et al., in press: fig. 11A); 1, long, ger than left one; 1, right paramere shorter than distinct, usually reaching posterior part of corium left one. In all Felisacus species state 0 is pres- (as in Namyatova et al., in press: fig. 12A). The state ent; in outgroup species state 1 is present. 0 is present in all species of Felisacus and Within Bryocorinae state 0 is also present in Nesidiocoris tenuis. Eccritotarsini. 20 Inner margin of cuneus shape: 0, straight (Namyatova 30 Apex of right paramere shape: 0, almost straight or et al., in press: fig. 13A, F); 1, convex (Namyatova slightly concave; 1, distinctly concave (Namyatova et al., in press: fig. 13E, G). In Felisacus and et al., in press: fig. 13A, R). The 1 is present in F. outgroup species both states are present. nigrescens and F. solomonicus only. 21 C-shaped marking on hemelytron occurrence: 0, 31 Apical part of right paramere: 0, indistinct (e.g., fig. absent; 1, present. C-shaped marking on hemelytron 11H, L, T, V); 1, distinct (e.g., fig. 11A, C, E, J). In is present in some Felisacus species. Felisacus both states are present; in outgroup spe- 22 C-shaped marking on corium shape: 0, both arms cies only state 0 occurs. reaching or almost reaching R+M; 1, with anterior 32 Apical part of right paramere length: 0, as long as or arm far not reaching R+M; 2, without anterior arm. longer than medial part (e.g., fig. 11A, C, J, P); 1, This character was not coded for those species distinctly shorter than medial part (e.g., figs. 11E, which do not have c-shaped marking (see char. 21). AB, 12G, I). This character was not coded for those State 1 is present in F. cristobalus and F. fedori, state species, where apical part indistinctly delimited (see 2 is present in F. curvatus and F. luzonus. char. 31). State 1 is present in F. australicus, F. 23 Tarsus shape: 0, straight (Namyatova et al., in press: fedori, F. javanus, F. lambkinae, F. luzonus, F. ochra- fig. 19F);1, incrassate apically (Namyatova et al., in ceus, and F. senaru. press: fig. 19A, C). State 1 is present in all Felisacus 33 Outer margin of right paramere shape: 0, straight, species and Pachypeltis reuteri. This state is present slightly concave or convex (e.g., fig. 11A, C, H); 1, in all Bryocorinae except of Dicyphinae (see Cassis, distinctly concave (e.g., figs. 11E, 12G, I);2, 1984; Namyatova et al., in press). rounded (e.g., figs. 11V, T, 12M). This character 24 Parempodia symmetry: 0, symmetrical (Namyatova was not coded for Nesidiocoris tenuis and Pachy- et al., in press: fig. 20D, F); 1, asymmetrical (Namy- peltis reuteri, in which right paramere is very atova et al., in press: fig. 20A, C). Among the spe- small. In Felisacus all states are present; in Stenotus cies in the analysis, parempodia are asymmetric in binotatus state 2 is present. Felisacus only. They are also asymmetrical in Eccri- 34 Inner margin of middle part of right paramere totarsini and some groups of Cylapainae (see shape: 0, without swelling, straight or concave Namyatova et al., in press). (e.g., fig. 11R, T, AB); 1, present (e.g., fig. 11A, C, Male genitalia E). This character was not coded for Nesidiocoris 25 Right side of genital capsule shape: 0, not folded tenuis and Pachypeltis reuteri, in which right (e.g., fig. 14A, B, E, X); 1, folded (e.g., fig. 14V, Y). paramere is very small. In Felisacus all states are State 1 is present in some species of Felisacus. present, in Stenotus binotatus state 0 occurs. 2016 NAMYATOVA AND CASSIS: FELISACUS DISTANT 167
35 Middle part of right paramere width: 0, subequal to 43 Outgrowth on posterior side of left paramere apically: width of basal part (e.g., fig. 11C, E, H); 1, twice as 0, absent; 1, present (e.g., fig. 11M, S, U, AC). In wide as basal part (e.g., figs. 11A, AF, 12A, C); 2, Felisacus both states are present; in outgroup spe- narrower than basal part (e.g., figs. 11L, AB, 12M, cies only state 0 is present. W). This character was not coded for Nesidiocoris 44 Tooth on posterior side of left paramere medially: tenuis and Pachypeltis reuteri, in which right para- 0, absent; 1, present (fig. 11G). Within Felisacus mere is very small. In Felisacus all states are present, both states are present; in outgroup this tooth is in Stenotus binotatus state 1 occurs. always absent. 36 Outer angle of right paramere occurrence: 0, absent 45 Apical part of left paramere shape: 0, not flattened; (e.g., figs. 11R, T, AB, 12M); 1, present (e.g., figs. 1, flattened (figs. 12F, X, 13H, J). State 1 is present 11A, C, W, 12U). only in F. luzonus, F. magnificus, F. philippinensis, 37 Inner angle of middle part of right paramere occur- and F. ponaponensis. rence: 0, indistinct or rounded (figs. 11A, E, N, 12A, 46 Outer margin of left paramere shape: 0, straight or C); 1, present, distinct (e.g., fig. 11R, T, AB). In Feli- convex; 1, distinctly concave (figs. 11U, AC, 12R, sacus and outgroup species both states occur. 13AE). State 1 is present in F. cristobalus, F. fedori, 38 Swelling or outgrowth at the inner margin of proxi- F. longiceps, and F. zuparkoi. mal inflection occurrence: 0, absent (e.g., figs. 11V, 47 Inner margin of left paramere shape: 0, without 12U, 13G); 1, present (e.g., figs. 11T, AB, 12M, W). swelling or outgrowth; 1, with single swelling or This character was not coded for those species, in outgrowth (e.g., figs. 11M, AC, S 13J); 2, with two which inner angle of middle part of right paramere or three swelling or outgrowths (figs. 12V, 13H). In indistinct or rounded (see char. 38). In Felisacus Felisacus all states are presentl in outgroup species both states are present. only state 0 occurs. 39 Outgrowth on inner angle of right paramere 48 Outgrowth on inner margin of left paramere length: 0, length: 0, short, in shape of swelling (figs. 11T, short, in shape of rounded swelling (figs. 11M, 12N); 13AB); 1, elongate (e.g., figs. 11AB, R, 12M). 1, short, triangular (figs. 11U, AC, 13AC, AE); 2, This character was not coded for those species, elongate (figs. 11S, 12R, X). This character was coded in which inner angle of middle part of right para- only for those species, where single swelling or out- mere indistinct or rounded, or present and not growth on inner margin of left paramere is present swollen (see chars. 37, 38). State 0 is present in F. (see char. 47). State 0 occurs in F. bismarckensis and cristobalus and F. yasunagai. F. lindbergae, state 1 is present in F. cristobalus, F. 40 Elongate outgrowth on inner angle of right paramere fedori, F. yasunagai, and F. zuparkoi; state 2 is present shape: 0, straight (figs. 11AB, 12M); 1, curved (figs. in F. ceylonicus, F. longiceps, and F. magnificus. 11R, 12Q, W, 12AD). This character was coded only 49 for those species, in which outgrowth on inner Teeth on outgrowth on inner margin of paramere angle of right paramere elongate (see char. 39). occurrence: 0, absent; 1, present (e.g., figs. 11U, 13J, State 0 is present in F. fedori and F. lindbergae. AE). This character was coded only for those spe- cies, where single swelling or outgrowth on inner 41 Basal part of right paramere relative length: 0, less margin of left paramere is present (see char. 47). than 0.3× as long as rest of paramere (e.g., fig. 11A, State 1 is present only in F. cristobalus, F. ponapo- C, E); 1, subequal to or longer than half of rest of nensis, and F. zuparkoi. the paramere (e.g., fig. 11N, R, T). This character was not coded for Nesidiocoris tenuis and Pachy- 50 Outgrowth on dorsal side of apical part of left para- peltis reuteri, in which right paramere is very small. mere occurrence: 0, absent; 1, present (e.g., fig. 11S, In Felisacus both states are present, in Stenotus U, AC). In Felisacus both states are present; in out- binotatus state 0 occurs. group species this outgrowth absent. 42 Setae on right paramere position: 0, placed on mid- 51 Setae on left paramere position: 0, placed mostly dle part along inner margin (e.g., fig. 11A, C, E); 1, near outer margin (e.g., fig. 11B, F, Q); 1, bunch of placed on inner angle (e.g., figs. 11T, AB, 12M, U). setae placed on inner margin (fig. 11S, U, AC). This This character was not coded for outgroup species, character was not coded for outgroup species, as as the position of setae is unclear. In Felisacus both well as for F. ponaponensis and F. yasunagai, where states are present. those setae were not observed. 168 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 403
52 Distal part of ductus seminis shape and sclerotiza- 62 Spicule I (convolute basally, placed close to middle tion: 0, entirely membranous; 1, only small apical line) occurrence: 0, absent; 1, present (fig. 9B, C). This part of ductus seminis sclerotized (e.g., figs. 8–9); 2, spicule is present only in F. meilingae and F. myersi. at least apical half of ductus seminis sclerotized 63 Spicule F (long, needle shaped, placed from left-hand (e.g., fig. 10C, D, F). In Felisacus states 1 and 2 are side) occurrence: 0, absent; 1, present (fig. 8A, J, K, present; in outgroup species 0 and 1 occur. L). This spicule is present in F. albus, F. glabratus, F. 53 Elongate sclerotized part of ductus seminis shape: 0, indicus, and F. insularis. straight apically (fig. 10A, I, L); 1, curved apically 64 Spicule G (long, wide and concave, placed from left- (e.g., fig. 10C, D, F). This character was coded only hand side) occurrence: 0, absent; 1, present (figs. 8E, for those species, in which at least apical half of I, 9B, C). This spicule is present only in F. bellus, F. ductus seminis sclerotized (see char. 52). State 0 is filicicola, F. meilingae, and F. myersi. present only in F. bismarckensis, F. luzonus, and F. ponaponensis. 65 Spicule H (hook shaped, middle-sized[[?]], placed from left-hand side) occurrence: 0, absent; 1, present 54 Spicule A (convolute, from right-hand side): 0, (figs. 8C, 9F). This spicule is present only in F. aus- absent; 1, present. In Felisacus both states are pres- tralicus and F. ochraceus. ent; in outgroup species this spicule is absent. 66 Distinct teeth on spicules or vesica occurrence: 0, 55 Spicule A shape: 0, curved (e.g., fig. 8C–F); 1, absent; 1, present (figs. 9G, N, 10B, E, M). State 1 is almost straight (fig. 9B, C). This state was coded present in F. bradi, F. ovalau, F. vitilevu, Nesiciodco- only for those species, in which spicule A is present ris tenuis, and Stenotus binotatus. (see char. 54). State 1 occurs only in F. meilingae and F. myersi. Female genitalia 56 Spicule B (curved right side): 0, absent; 1, present 67 Dorsal labiate plate size: 0, wider than distance (e.g., fig. 8A–C). In Felisacus both states are present; between valvula apodemes (fig. 16C, I, L, N); 1, in outgroup species this spicule is absent. very small, entirely placed between valvula apodemes (fig. 16A, E, G, K). In Felisacus both 57 Apical arm of spicule B shape: 0, almost straight or states are present; in outgroup species only state 0 slightly curved and not very long (e.g., fig. 8A, B, E, is present. G); 1, present, long (figs. 8C, F, 9F). This state was coded only for those species, in which spicule B is 68 Semioval sclerite on dorsal labiate plate occurrence: 0, present (see char. 56). State 1 is present only in F. absent; 1, present (fig. 16C, I). In Felisacus both states australicus, F. caledonicus, and F. ochraceus. are present; in outgroup species only state 0 occurs. 58 Spicules C (convolute from left-hand side) occur- 69 Sclerotized rings on dorsal labiate plate occurrence: rence: 0, absent; 1, present (e.g., fig. 8B–D). InFeli - 0, absent (fig. 16A, E, G, K, N); 1, present (fig. 16C, sacus both states are present; in outgroup species I). In Felisacus both states are present; in outgroup this spicule is absent. species only state 1 occurs. 59 Spicule C base position: 0, approximately at the 70 Striation on dorsal labiate plate occurrence: 0, same level with other spicules (e.g., fig. 9A–C); 1, absent; 1, present (fig. 16C, L, N). State 1 is present distinctly moved distally (figs. 8B, D, P, 9I). This only in F. bradi, F. curvatus and F. wangae. state was coded only for those species, in which 71 Spermathecal gland position: 0, placed medially or spicule B is present (see char. 58). State 1 is present posteriorly of dorsal labiate plate (fig. 16I, L, N); 1, in F. amboinae, F. bau, F. linae, and F. senaru. placed at the anterior margin of dorsal labiate plate 60 Spicule D (small, placed at base) occurrence: 0, (fig. 16A, E, G). In Felisacus both states are present; absent; 1, present (fig. 9B, C). This spicule is present in outgroup species only state 0 occurs. only in F. meilingae and F. myersi. 61 Spicule E (weakly sclerotized, from right-hand side) The associated Data Matrix can be viewed at occurrence: 0, absent; 1, present (figs. 8I, R, 9M). http://dx.doi.org/10.5531/sd.sp.22 as a .nex file, a State 1 is present in F. filicicola, F. lorhowensis, and format currently supported by the Mesquite F. tanna. Project. Scientific Publications of the American Museum of Natural History American Museum Novitates Bulletin of the American Museum of Natural History Anthropological Papers of the American Museum of Natural History
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On the cover: Male specimen of Felisacus senaru, new species.