The Complete Mitochondrial Genome and Novel Gene Arrangement of the Unique-Headed Bug Stenopirates sp. (Hemiptera: Enicocephalidae)

Many of true bugs are important insect pests to cultivated crops and some are important vectors of human diseases, but few cladistic analyses have addressed relationships among the seven infraorders of Heteroptera. The Enicocephalomorpha and Nepomorpha are consider the basal groups of Heteroptera, but the basal-most lineage remains unresolved. Here we report the mitochondrial genome of the unique-headed bug Stenopirates sp., the first mitochondrial genome sequenced from Enicocephalomorpha. The Stenopirates sp. mitochondrial genome is a typical circular DNA molecule of 15, 384 bp in length, and contains 37 genes and a large non-coding fragment. The gene order differs substantially from other known insect mitochondrial genomes, with rearrangements of both tRNA genes and protein-coding genes. The overall AT content (82.5%) of Stenopirates sp. is the highest among all the known heteropteran mitochondrial genomes. The strand bias is consistent with other true bugs with negative GC-skew and positive AT-skew for the J-strand. The heteropteran mitochondrial atp8 exhibits the highest evolutionary rate, whereas cox1 appears to have the lowest rate. Furthermore, a negative correlation was observed between the variation of nucleotide substitutions and the GC content of each protein-coding gene. A microsatellite was identified in the putative control region. Finally, phylogenetic reconstruction suggests that Enicocephalomorpha is the sister group to all the remaining Heteroptera

Taxonomy based on science is necessary for global conservation

Peer reviewe

A new Xenicocephalus species from Ecuador (Heteroptera, Enicocephalomorpha, Enicocephalidae)

Xenicocephalus tomhenryi sp. n. (Insecta: Hemiptera: Heteroptera: Enicocephalomorpha: Enicocephalidae) is established for a single macropterous female from Ecuador. The enigmatic genus now includes three species known from only two Neotropical adults and an incomplete female specimen. The new species is described and illustrated, extensive comparative diagnoses for Xenicocephalus species are provided, and nomenclature, distribution, and biology of the genus are reviewed. The architecture of the raptorial forelegs of Xenicocephalus is unique among Enicocephalomorpha, and the genus is classified as subfamily incertae sedis

Pseudohenschiella hauseri Baňař & Štys 2006, sp. nov.

Pseudohenschiella hauseri sp. nov. (Figs. 1-13) Type locality. Madagascar, Antananarivo prov., Ankaratra hills, Manjakatompo Forest Station, 1980 m a.s.l. Type material. HOLOTYPE: ♀, ‘Mad-89/23: Madagascar (Prov. / Antananarivo, sous-préf. / Ambatolampy): Massif Ankaratra, / Station Forestière Manjakatompo, / près du sommet du Anosiarivo, foręt / primaire, sous écorces, 1980m; / 26.xi. 1989; leg. B. Hauser’. The specimen bears the red type label ‘ HOLOTYPE / Pseudohenschiella / hauseri sp.nov. / Baňař & Štys det. 2006’. Holotype slightly damaged (midtibiae partially broken), card-mounted, right foreleg mounted separately on another card; to be deposited in the collection of Muséum d’Histoire Naturelle, Geneva (Switzerland). Description. Measurements (all in mm; L = length, W = width, max. = maximum, min. = minimum). Total L – 2.75; head: anterior lobe L – 0.26, posterior lobe L – 0.18, posterior lobe W – 0.25, distance of eye to apex of antennifer – 0.13, diatone (max. W across eyes) – 0.24, min. interocular distance, dorsal – 0.15, min. interocular distance, ventral – 0.11, eye L – 0.08; labium: total L – 0.44; antenna: segment 1 L – 0.11, segment 2 L – 0.24, segment 3 L – 0.22, segment 4 L – 0.21; pronotum: total L (max.) – 0.46, collum L (median) – 0.13, midlobe L (max.) – 0.19, hindlobe L (max.) – 0.14, collum max. W – 0.26, midlobe, max. W – 0.46, hindlobe, max. W – 0.53; foreleg: forefemur L – 0.40, foretibia L – 0.33, forefemur max. W – 0.18, foretibia max. W – 0.16; midleg: midfemur L – 0.35, midtibia L – 0.30, midtarsus L (without claw) – 0.11, midfemur max. W – 0.10, midtibia max. W – 0.07, midtarsus max. W – 0.05; hindleg: hindfemur L – 0.42, hindtibia L – 0.48, hindtarsus L (without claw) – 0.16, hindfemur max. W – 0.13, hindtibia max. W – 0.08, hindtarsus max. W – 0.03. Coloration (Fig. 1). Head and thorax brown; forewing membrane slightly paler, extreme apex of head, labium, forewing veins, legs and antennal segments 1-3 yellowish brown, posterior (inner) faces of femora somewhat infuscate; antennal segment 4 whitish; dorsum and venter of abdomen beige (small isolated sclerites brown). Cuticle. Head, thorax and abdomen generally matt, with clearly visible punctation and minute setigerous tubercles; legs (with some exceptions – see foreleg structures below), antennae, labium and small median impression on midlobe of pronotum smooth and lustrous. Head, thorax and extremities strongly sclerotized, abdomen weakly so. Vestiture whitish, relatively dense. Setae on dorsal face of head prominent, curved anterad, becoming longer towards antennifers. Lateral faces nearly bare. Setae on ventral face of head (Fig. 2) strongly prominent, longer than setae on dorsal and ventral faces; on anterior lobe curved posterad, on posterior lobe curved anterad (excepting the basal part, adjoining the collum). Collum covered by dense hairs curved irregularly in different directions. Dorsal and lateral parts of mid- and hindlobe of pronotum and ‘pleural’ parts of thorax covered with dense prominent hairs, these becoming longer ventrally; setae of both dorsum and venter of thorax directed caudad. Abdomen with adpressed hairs, laterotergites 7-9 with several very long, trichobothria-like setae, sternum 7 and subgenital plate covered with long, dense, semi-erect setae. Forewing veins pilose, cells bare, very rarely with one seta per cell. Wing margin of four distinct sectors differing in pilosity, as follows: (a) proximal half of anterior margin with one row of uniform, short, curved setae; (b) third quarter of anterior margin with same hairs as (a) but complemented by second row of longer, curved hairs, both rows becoming longer distad; (c) beginning of distal quarter of anterior margin with admixed third row of erect bristles, the latter shortly disappearing and the rest of anterior margin, whole apical margin, and distal part of posterior margin (up to entry of last remigial vein) formed by one row of slightly curved, diagonal, alternating short and very long hairs; (d) whole proximal part of posterior margin bare. Antennae. All segments with rather long diagonal setae; distal part of segment 2 and whole segments 3 and 4 also with more erect, subvertical, long, fine setae about twice as long as segment diameter. Labium with dense, semi-erect, slightly curved pilosity, particularly long and conspicuous at dorsal surface of segment 3. Forelegs with two types of setae: (a) long, prominent trichobothria-like setae (tr-setae); (b) shorter, semi-erect, denser setae. Coxa and trochanter antero- and posteroventrally with numerous (ca 20-30) semi-erect setae and several (3-5) tr-setae. Dorsal face of trochanter with rows of five setae, one longest, other four subequal in length. Femur and tibia covered more densely and regularly on all surface, with exception of small bare area on antero-basal part of femur. Femur with 15-20 regularly distributed tr-setae with exception of anterior face. Tibia with 10-12 tr-setae, distributed especially on posterodorsal face, tarsus with 4-5 tr-setae. Mid- and hindlegs. Both the mid- and hindfemur with a conspicuous, outstanding, curved and long setae in two thirds of dorsal face, ventral face with shorter, adpressed setae. Mid- and hindtibiae and tarsi with dense, semi-erect setae on all faces. Head (Fig. 2). Slightly shorter than pronotum (ratio 0.95). Anterior lobe cylindrical, longer than posterior lobe (ratio 1. 44). Postocular constriction deep. Posterior lobe transverse, lateral margins irregularly rounded, with slightly visible median groove, widest in basal third, ratio length to maximum width 0.72. Ratio length of eye to distance eye – apex of antennifer 0.62. Eyes medium-sized, dark reddish brown. Ocelli on small tubercles, directed anterolaterad, each with narrow carmine ringlet on base. Dorsal ocular index 5.3, ventral ocular index 3.7. Neck very short. Antennae (Fig. 1) relatively short and thin, shorter than head and pronotum together, antennal formula (longest segment first): 2-3-4-1 (length differences 2, 3, 4 very small). Segments 2 and 3 moderately subfusiform, 4 fusiform. Labium (Fig. 2) very short, thick, directed anterad, labial formula (longest segment first) 3-4-2-1, segment 1 minute, 3 basally constricted, and with medially inflated ventral side. Pronotum (Fig. 1). Collum thick, simple, without lateral tubercles, precollum narrow, transverse impression between collum and midlobe strikingly deep and wide, broadly V-shaped. Lateral margins of midlobe rounded, its median marked by broad, inversely T-shaped impression (its stalk shallow, interrupting anterior margin; its transverse bar deep, rounded, distant from posterior margin), disc with a pair of deep anterolateral pits; lateral and medial parts of posterior margin rounded, sublateral ones shallowly concave. Hindlobe moderately wider than midlobe, widest in middle, lateral sides very moderately rounded, posterolateral angles roundedly subrectangular, posterior margin shallowly concave. Mesoscutellum large, triangular, disc elevated with anterior row of 2+2 muscle impressions and unpaired posterior impression, apex produced in rounded mucro. Fore acetabula open, ‘proepimeral lobe’ (cum posterior prosupracoxale) reaching half width of forecoxa. Terminalia. Segment 8 clearly subdivided into fully sclerotized tergum, incompletely sclerotized laterotergites (occupying truly lateral position), and ventral, sclerotized subgenital plate. Posteromedial margin of the latter emarginate, the space filled by intersegmental membrane 8-9 provided with small medial sclerite. Segment 9 ring-shaped, fully sclerotized, its ventral (subproctigeral) part narrow. Proctiger globular. Differential diagnosis. Pseudohenschiella hauseri sp. nov. differs from P. minuscula Villiers, 1969 by longer body, wider foretibia, and shape of posterior lobe of head and collum (very short in P. minuscula); from P. usingeri and P. flavipes Villiers, 1969 by wider foretibia, different dimensions and shape of both mid- and hind- pronotal lobes, and non-opposite position of transverse veins associated with forewing discal cell; from P. brevipes Villiers, 1958 by longer collum, different shape and proportions of posterior lobe of head, and by different shape of forewing discal cell (see the Key, and VILLIERS 1969). Dimensions of fore tibia may be sexually dimorphic (tibia distally broader in females); so far all species are only known from single individuals. Etymology. Named after Bernd Hauser (Geneva), collector of the species. Bionomy. The holotype was collected under bark in a primary forest.Published as part of Baňař, Petr & Štys, Pavel, 2006, A new species of Pseudohenschiella (Heteroptera: Enicocephalidae) from Madagascar, pp. 7-14 in Acta Entomologica Musei Nationalis Pragae 46 on pages 8-13, DOI: 10.5281/zenodo.450353

The first species of Systelloderes (Hemiptera: Heteroptera: Enicocephalidae) from New Caledonia

Štys, Pavel, Baňař, Petr (2007): The first species of Systelloderes (Hemiptera: Heteroptera: Enicocephalidae) from New Caledonia. Acta Entomologica Musei Nationalis Pragae 47: 3-15, DOI: http://doi.org/10.5281/zenodo.450356

Enicocephalus flavicollis Westwood 1837

Enicocephalus flavicollis Westwood, 1837 - male, redescription (Figs. 1 –6, 10– 17) Material examined. Lectotype: male, first label: // I. S ti. Vincentii // [handwritten = Insula Sancti Vincentii (Latin)]; second label: // Enicocephalus / flavicollis Westw. // [handwritten]; third label: // Type Hem: 4 / 6 / Enicocephalus / flavicollis / Westwood / Hope Dept. Oxford // [partly printed and partly handwritten]. Specimen bears red label: // Enicocephalus flavicollis / Westwood, 1837 / LECTOTYPE ♂ / P. Štys & P. B aňař det. 2011 // [printed]. Specimen dry mounted, glued to triangle card, in very bad condition: midlobe of pronotum (and some sternal parts) strongly damaged, antennal segments III, IV, left mid- and hindtibiae and tarsi are missing, left hindfemur mounted separately (Fig. 1). Deposited in Hope’s collection, University Museum of Natural History, Oxford. Paralectotype: male, first label: // Windward side / St. Vincent, W.I. / H.H. Smith 190 // [printed; number ‘ 190 ’ handwritten]; second label: // 95-206. //; third label: // H enicocephalus [originally Enicocephalus - overwritten] / flavicollis Westw. // [handwritten]; specimen bears red label: // Enicocephalus flavicollis / Westwood, 1837 / PARALECTOTYPE ♂ / P. Št ys & P. B aňař det. 2011 // [printed]. Dry mounted specimen glued to a triangle, in good condition, right antennal segment 4 and left hind leg missing. Deposited in Department of Entomology, Natural History Museum, London. Measurements (in mm): lectotype first (paralectotype in parentheses, italics). Total body length —ca 3.9 (specimen damaged) (3.87). Head (without neck). Total length— 0.62 (0.63); posterior lobe, length— 0.24 (0.23), width— 0.38 (0.37); distance of eye to apex of antennifer— 0.13 (0.13); diatone (maximum width across eyes)— 0.39 (0.38); dorsal synthlipsis (minimum interocular distance)— 0.18 (0.19); eye, length— 0.17 (0.16); gena, length— 0.03 (0.03), width (minimum)— 0.19 (0.18). Labium. Total length— 0.47 (0.46); segment I, length— 0.06 (0.05); segment II, length— 0.07 (0.07); segment III, length— 0.20 (0.20); segment IV, length— 0.13 (0.13). Antenna. Segment I, length— 0.14 (0.13); segment II, length— 0.29 (0.28), basal width— 0.02 (0.02), distal width— 0.05 (0.05); segment III, length— 0.37 (0.37); segment IV, length— 0.36 (0.36). Pronotum. Total length (median)—ca 0.58–0.62 (specimen damaged) (0.60); collum, length (median)— 0.13 (0.13), width (maximum)— 0.29 (0.29); midlobe, length (median)—ca 0.25–0.29 (specimen damaged) (0.27), width (maximum)— 0.54 (0.55); hindlobe, length (maximum)— 0.40 (0.38), length (mediane)— 0.19 (0.18), width (maximum)— 0.80 (0.78). Foreleg. Femur, length— 0.82 (0.81), width (maximum)— 0.20 (0.19); tibia, length— 0.74 (0.72), width (maximum)— 0.17 (0.17). Coloration (Figs. 1, 3, 5). Bicolorous, piceous or yellow, the colours strongly contrasting except on abdomen. No red or scarlet pigments. Head and thorax. Dorsum. Dark body parts piceous brown (dorsum of head, most of the posterior lobe of pronotum, mesoscutellum); neck of the head somewhat lighter; non-melanized parts unicolorous, contrastingly yellow (collum, midlobe of pronotum; linear, transverse, short, anterior strip of the hindlobe of pronotum). The yellow transverse strip of posterior lobe of pronotum with 1 + 1 submedial extensions shaped as segments of circle. Lateral sides and venter of head and thorax. Anterior lobe of head (from apex to postocular constriction) piceous, posterior lobe yellow; prothorax yellow (the yellow area confluent with the anterior yellow strip on posterior lobe of pronotum), pterothorax (incl. all supracoxal lobes) piceous, pterosterna yellowish. Antennae piceous brown; proximal part of segment IV non-contrastingly lighter. Labium: segments I and II brownish, III and IV yellow; legs (inclusive coxae) uniformly yellow. Forewings piceous brown with blackish hue (Figs. 1, 3), veins concolorous except for the noncontrastingly yellowish marginal vein (costal margin cum ambient vein up to the level of cu-an). The light coloration of the marginal vein invisible in transmitted light owing to semitransparency of the wing; ventral side matching the dorsal one. Hindwings blackish-brown. Abdomen. Dorsum not examined; venter: segments 1–3 yellowish brown, 3–7 piceous, 8 and 9 yellowish brown; posteromedial part of 2 and medial parts of 3–5 black. Coloration of paralectotype London male matching the redescription of Oxford lectotype male (Wyhodzinsky& Schmidt 1991) except for the lectotype abdomen said to be whitish. Microsculpture. Cuticle strongly lustrous and smooth. No setigerous tubercles. Posterior lobe of pronotum irregularly shallowly rugulose. Ventral surface of fore trochanter with about 5 transverse rows of serrate microstructures. Ventral face of forefemur with a continuous percurrent longitudinal strips of minute black granules; these terminating nearly apically, leaving the area contacting basis of foretibia bare but provided with one large posterior dens; no microtrichiae or spicules present. Venter of abdomen with minute transverse wrinkles. Vestiture. Macrotrichia (further on ‘setae’) golden, rather soft, usually short and straight, rarely curved. Head. Dorsal view. Anterior lobe of head covered by short and appressed, very dense short setae, obscuring its surface; longer curved setae occurring apically and in front of eyes only. Posterior lobe with sparse straight to curved, rather short setae. Lateral view. Dorsum with long, very dense, regularly curved, diagonal setae directed anteriad (excl. on preocular lobe), those on posterior lobe obscuring its outline. Venter with long, anteriorly directed, long straight or curved setae forming a conspicuous barb on the posterior lobe. Ventr al view. Very long and irregular macrotrichial cover all over. Eyes with a few straight interommatidial setae in anterior half, with many curved setae in posterior half. Antennae with short, dense, diagonal pubescence; segment 2 with few semi-erect hairs on its inner side, segment 3 on both sides and apically, segment 4 all over. Labium. Segment 3 with moderately long and moderately dense diagonal straight macrotrichia interspersed with long, thin, erect to suberect trichobothrium-like setae along both sides (4 + 4) and clustering in the apical part. Thorax. Dorsal view. Lateral outline: collum with moderately dense, straight short setae, midlobe with curved setae gradually longer caudad, hindlobe with very dense, curved, short setae. Mesoscutellum with long curved dense setae. Lateral view. Dorsum densely covered by extremely short, and very dense cover of straight to curved, diagonal, minute setae, only those on collum with a few erect setae. Prothoracic presupracoxale with a cluster of strikingly long hairs. Forewings with short, curved hairs (mixture of golden and black ones) distributed in a manner modal for Enicocephalus (cf. Wygodzinsky & Schmidt 1991). Intervenial membrane bare. Hindwings bare. Legs. Anterior faces of all coxae densely pilose. Foreleg. Dorsal face of femur with long diagonal setae, with interspersed few trichobothrium-like diagonal setae (recognizable only by their thinness); ventral face with long, thin, straight setae. Tibia covered all over by short, medium long, mostly appressed to diagonal straight setae (similar to those on dorsal face of femur), occurring on the anterior longitudinal tibial depression as well; dorsal and ventral faces additionally with numerous trichobothrium-like setae, all thin and all erect, rarely straight, mostly curved at the tip, nearly as long as local tibial diameter. The latter kind of setae occurring all over tarsus. Mid - and hindtibiae with a long diagonal vestiture all over, setae longer than tibial diameters; the same true for mid - and hindtarsi. The setae of dorsal and ventral tibial faces thin, trichobothrium-like (?). Abdomen. Margin without specialized setae, uniformly covered with uniformly curved setae becoming gradually longer posteriad, longest all along edge of segments 7 and particularly 8; the latter posteriorly with 3 + 3 very long setae, the longest exceeding apex of pygophore. Venter. All segments (exc. pygophore) densely covered with very short and uniform diagonal pubescence with setae slightly dilated (not gradually pointed) at tips, a few longer setae present at the medial keel only. Structure. Very small species, but rather robustly built. Head. Dorsal view (Figs. 1 –2, 6, 11). Anterior lobe short, insertion of antennae far from apex of anteclypeus, midway beween eyes and anteclypeus, gena very short, ratio gena lenght to its width 0.17; ratio of gena length to length of eye 0.19. Postocular constriction deep and long (about as long as the gena) and sharply delimited. Postocular lobe strikingly transverse, laterally asymmetrically rounded, widest behind the middle, ratio of its length to maximum width 0.62, no structural indication of the median except for a slight interocellar concavity. Ocelli situated on low tubercles, interocellar distance shorter than distance eye–ocellus. Head. Lateral view. Dorsal outline of the anteocular part of anterior lobe straight, slanting ventrad, strongly convex over the eye. Posterior lobe strongly convex, bulbous, its outline much exceeding that of the anterior lobe (more than shown by Wygodzinsky & Schmidt (1991, Fig. 80 B)). Ventral outline of preocular part of anterior lobe straight, that of posterior lobe moderately convex. Eyes large, strongly convex and outstanding, inner margins convex in dorsal view, ommatidia separately convex. Dorsal ocular index 2.00. Lateral view: dorsal margin far distant from dorsal margin of head, ventral margin slightly exceeding ventral margin of head. Ventral view: eyes strongly approaching each other. Antennae short, first segment unusually thick, 2 nd subclavate, strongly widening towards apex, segments III and IV stick-shaped, much thinner than segment II (but not flagelliform). Antennal formula (longest segment first) III-II-IV-I. Labium. Segments I and II very short, their dorsal faces flat as well as that of segment III; the latter rather short, conical. Labial formula (longest segment first) III-IV-II-I. Thorax. Pronotum in dorsal view . (Figs. 1 –2, 6, 11). Collum long and laterally rounded, lacking lateroventral tubercles and any structurally indicated median, but with an unusual feature (natural or artifact?): posterior 3 / 4 with one medial and 1 + 1 sublateral subangular shallow impressions, contacting each other and reaching the short and sharp constriction between collum and midlobe. Midlobe. Dorsal view. (Owing to the light colour and strong lustre of the area the description of surface sculpture is open to reinterpretation). Lateral margins strongly rounded, level of maximum width close to the anterior margin, posterior margin nearly straight, not interrupted. Ratio of midlobe maximum width to median length 3.05. Medial impression: antero-medial sector slightly depressed, with two impressed lines meeting in the centre and delimiting an anteromedial acutangular triangle with an apex pointing centrad; apex of the triangle sloping into a deep pit with a more shallow lateral linear extensions and rampart-like posterior margin coinciding with the posterior margin of the lobe itself (the impression resembling Oncylocotis -like inverse T-shaped impression.). The impression containing a linear, impressed median. Sublateral impressions present as simple, shallow, depressions, not interrupting the posterior margin of the midlobe and situated anterolaterad to the median depression. Posterior lobe. Dorsal view. Lateral margins regularly rounded inclusive indistinctly indicated posterolateral angles; medial part of posterior margin deeply angularly excised, lateral parts of this excision convex. No median structure. Ratio of hindlobe maximum width to its maximum length 2.05. Pronotum in lateral view. Collum with no ventrolateral tubercles. Lateral sides of pronotum not clearly delimited from supracoxal lobes, no horizontal lateral ridge or sulcus present. ‘Proepimeral lobe’ short, rounded, ventrally not reaching the level of apex of posterior prosupracoxale. Mesoscutellum with a convex surface, short, strongly rounded lateral margins, and unusually broad and long, apically rounded mucro separated from the disc by a transverse impression. Wings macropterous, not caducous. Forewings much exceeding the abdomen, venation (Fig. 13) as illustrated by Wygodzinsky & Schmidt (1991, Fig. 80 C), only the distal sector of AA + AP continuous, complete (checked on both the specimens). Hindwings longer than abdomen Legs. Foreleg. For microsculpture of trochanter and femur see above. Dorsal face of distal part of trochanter convex, slightly elevated over the neighbouring part of femur; ventral basis of femur slightly angularly produced and exceeding the surface of trochanter ventrad; ratio of femur length to maximum width 4.26. Tibia abruptly widening on dorsoventral plane in about a third of its length; ratio of tibia length to maximum width 4.24. Longitudinal (anterior) tibial depression starting at the points of tibial dilatation and running up to the cleaning comb. (Foretarsus pressed to the distal edge of tibia on both forelegs - consequently, apicitibial and tarsal armatures could not have been studied). Inner (anterior) claw large, posterior one represented by a short rounded hump only. Mid - and hindtibiae with two apical bristle combs each; claws nearly isomorphic (the inner (anterior one) slightly longer. Abdomen. Venter sclerotized, sulci between ventral laterotergites and sterna marked by deep impression and vaguely delimited laterosternal plates. Segments 3–7 with a sharp median keel, segment 8 without the keel, simply trapezoidal, posterolateral angles not protruding, embracing strikingly large, broadly transverse pygophore. Pygophore (Figs. 15–16) (as seen in strictly posterior view and examined under 100 magnification; dorsum covered by abdominal segment 8 and wings) strongly transverse, posterior face oval, delimited by a rampart-like swollen margin of the posterior foramen of the pygophore. Guide slender and small, of a modal shape, with distinct shaft, head and frame, distal margin of guide head truncate. Supradistal plate (Fig. 16) seen as a sclerite dorsad and anterad to guide, parameres situated laterad to the former. Diagnostic characters. Small species (3.9 mm), contrastingly bicolorous (lacking red pigment, yellow and brown - shared with E. boraceianus, E. guarani, E. schuhi (length 6.5–6.7 mm), and E. tupi only), legs uniformly yellow, collum and midlobe of pronotum uniformly yellow, the yellow part of pronotum involving also a short stripe on the disc of the hindlobe (unique!). Discal cell of forewing strikingly long and pointed, reaching nearly the wing margin (unique!). Male forelegs strikingly slender and genitalia small. Outer fore claw extremely reduced (shared with E. semirufus and E. usingeri only). Endemic to St. Vincent Island, sympatric occurrence of any other Enicocephalus species unknown.Published as part of Štys, Pavel & Baňař, Petr, 2012, Taxonomy and nomenclature of Enicocephalus flavicollis Westwood — type species of the type genus of the Enicocephalidae (Hemiptera: Heteroptera: Enicocephalomorpha), pp. 53-68 in Zootaxa 3237 on pages 56-59, DOI: 10.5281/zenodo.21417

Systelloderes loebli Štys & Baňař 2007, sp. nov.

Systelloderes loebli sp. nov. (Figs. 1-16) Type locality. New Caledonia, Mount Koghi, 400-500 m a.s.l. Type material. HOLOTYPE: ♀, ‘ New Caledonia / Mt. Koghi, prim. for. / 400-500m, 18.-19.x / 1998, I. Löbl, litter’. The specimen bears the following red label: ‘ HOLOTYPE / Systelloderes / loebli sp. nov. / Štys & Baňař det. 2007’; collection of Muséum d’Histoire Naturelle, Genève (Switzerland). Diagnosis. Large species, over 5 mm long, macropterous, brownish. Antennal segment 2 longer than segment 3, segments 2-4 isomorphic, terete. Fore leg: coxa with a long, narrow, free, erect scale; femur with a prominent basidorsal extension rising high above the level of tarsus; femoro-tibial membrane with dorsal neopatellar sclerites; tibial process absent; tibial and tarsal armatures as illustrated. Description. Body elongate, moderately robust, extremities short, coloration and vestiture uniform and inconspicuous (Fig. 1). Measurements. Total body length – 5.75-5.85 (abdomen deformed). Head. Anterior lobe, L – 0.71; posterior lobe, L – 0.31, W – 0.38; distance of eye to apex of antennifer – 0.49; diatone (max W across eyes) – 0.36; min interocular distance, dorsal – 0.24; min interocular distance, ventral – 0.18; eye, L – 0.15. Labium. Total, L – 0.95; segment 1, L – 0.12; segment 2, L – 0.21; segment 3, L – 0.48; segment 4, L – 0.16. Antenna. Segment 1, L – 0.24; segment 2, L – 0.62; segment 3, L – 0.56; segment 4, L – 0.42. Pronotum. Total L (max) – 0.98; collum: L (median) – 0.18, max W – 0.49; midlobe: L (max) – 0.44, W (max) – 0.89; hindlobe: L (max) – 0.36, L (median) – 0.31, W (max) – 0.98. Forewing. Max L – 2.95. Fore leg. Total L – 1.07, max W – 0.42; Ti 1 : L – 0.87, max W – 0.41;. Ta 1 : L – 0.29, max W – 0.17; anterior claw, L (basis – apex) – 0.31; posterior claw, L (basis – apex) – 0.26. Middle leg. F 2 : L – 0.84, max W – 0.18; Ti 2 : L – 0.69, max W – 0.13; Ta 2 : L (without claw) – 0.24, max W – 0.09. Hind leg. F 3 : L – 1.02, max W – 0.29; Ti 3 : L – 1.16, max W – 0.14; Ta 3 : L (without claw) – 0.38, max W – 0.09. Coloration (Fig.1). Body nearly unicolorous, between light-brown (antennae, middle and hind legs) and brown to dark-brown (head and midlobe of pronotum). Texture. Moderately shiny (including extremities), head lustrous. Dorsum of head with irregularly distributed, rather sparse, small setigerous tubercles, lateral and ventral sides of head, prothoracic collum, and dorsum of pronotal midlobe with a continuous cover of regularly distributed, minute setigerous tubercles (the latter particularly distinct in preocular area, on posterior cephalic lobe and on midlobe of pronotum, creating their shagreened appearance). Setigerous tubercles replaced by indistinct, large and shallow alveoles laterally on parts of prothorax and dorsum of pronotal hindlobe, the latter slightly rugulose. Mesoscutellum, wings and abdomen without particular structures. Antennae, labium, Cx 1 , Tr 1 , Ta 1 , middle legs, and hind legs smooth. Anterior and posterior faces of F 1 and Ti 1 with scattered minute setigerous tubercles; longitudinal depression of anterior (mesal) face of Ti 1 with transverse wrinkles. Vestiture. Macrotrichia golden, ‘soft’, mostly short, straight, oblique, on some body parts (particularly posterior lobe of head, pronotum, and forewings) moderately to strongly curved. Macrotrichia mixed with much longer, diagonal, semierect to erect, conspicuous, mostly straight to only slightly curved trl setae with bilaterally symmetrical position; some of these may represent true trichobothria although the bothrium itself was not observed. Scales absent, except on Cx 1 (see below). Head, labrum, and labium. Distribution of trl setae (longitudinally arranged if more than 1+1) as follows. Head: several trl setae on and alongside anteclypeus and on labrum; 2+2 preocular (proximad to antennifers); 1+1 close to mesal eye margins, 2+2 postocellar; labial segment 1: dorsum/venter 1+1/0; 2: 3+3/0; 3: 3+3/2+2; 4: 3+3/3+3 (all strongly oblique and curved, similar to normal, elongate macrotrichia). Setation of ventral surface of head erect, short; longer and curved hairs on buccular part, gradually more curved and longer on proximal part close to neck, none trichobothrium-like. Antennae with uniform, short, straight, oblique setae; trl setae 1+1 on segment 1, then occurring from base of segment 2 to apex of segment 4, more elongate, denser and more erect distally. Prothorax. Dense, ‘soft’, short, mostly curved setae; trl setae: collum 1+1 anterolaterally, 1+1 posteromedially; midlobe: 1+1 posteromedially. Mesoscutellum. trl setae 2+2, in proximal part. Forewings with sparse, short, curved microtrichia only on veins, none on wing membrane. Fore leg. Cx 1 with curved, nearly adpressed pubescence and two long, distiventral, diagonally pointing trl setae; a long, narrow, moderately ectally curved scale (L 0.12, W 0.02) subapically on ventral surface, nearer mesal face (Figs. 3, 6). Tr 1 with curved, long setae; on ventral face, with 3-4 long, curved trl setae: the most distal one inserted at identical point as coxal scale; a few setae directed proximad, the other distad. F 1 with short, oblique setae on dorsal and ventral faces, anterior and posterior faces nearly bare; erect trl setae as follows: about 5+ 5 in double-row in distal half of dorsal face and multiple terminal dorsal cluster, and about 5+5 trl setae regularly distributed on ventral face. Ti 1 anterior face nearly bare, posterior and ventral faces with long, straight, oblique setae (particularly in distal half), dorsal and ventral faces and distal edge with numerous, irregularly distributed trl setae; other conspicuous macrotrichia: a single subpatellar trl seta, and three very long, curly setae at distiventral angle. Ta 1 with long, straight, diagonal setae, especially dense and mostly curly on ventral face; conspicuous trl setae: 1+1 dorsal, subterminal, strikingly long, and 1+1 apicilateral claw-guarding setae (anterior one, close to the shorter claw, conspicuously shorter). Middle and hind legs densely covered with short oblique to semierect setae on all faces; tibial and tarsal setae radiating relative to tibial and tarsal axes. Tr 2 and Tr 3 with two conspicuous erect ventral trl setae each. Ventral faces of F 2 and F 3 each with one basal (adtrochanteral), one postmedial, and several distal suberect to erect trl setae, dorsal faces of F 2 and F 3 each with 1+1 erect subterminal (adtibial) trl setae. Ti 2 and Ti 3 with long, oblique to erect trl setae developed along entire length,>15 and> 20 in number, respectively. Ta 2 and Ta 3 (second ’segments’) with about 10 strongly oblique trl setae on Ta 2 , over 12 on Ta 3 ; setae on the latter undoubtedly homologous but not thin and trl like, and nearly spiniform instead. The only visually undoubtedly trl seta on Ta 3 is a ventral erect seta on its first ‘segment’. Abdomen. Dorsum with short setae, without scales. Venter with golden short to moderately long, straight to slightly curved semierect macrotrichia (length and density increasing distally) intermixed with short, blackish, adpressed hairs. Some trl setae occurring marginally; distribution of long trl setae on posterior segments as follows: ventrite 7 – 1+1 at posterolateral angles, 1+1 submedial near posterior margin; subgenital plate: 1+1 submedial (extremely long) in basal part, 1+1 (very long) at posterior margin. Structure. Head (Figs. 1-2) strikingly narrow and long, slightly longer than pronotum (1.04 times as long as pronotum). Anterior lobe markedly longer than posterior lobe, 2.3 times as long. Lateral margin of preocular region parallel-sided proximally, its long distal part slightly convex, diverging towards antennifers; anteclypeus very long, narrow. Eye 0.31 times as long as distance between eye and antennifer. Postocular impression broad, shallow; lateral margin of postocular part of anterior cephalic lobe straight, diverging towards convex side of posterior lobe – postocular impression not marked in lateral outline. Posterior lobe transverse, 0.82 times as long as wide, its dorsum strongly convex, lateral sides moderately so. Eyes small, in lateral view not exceeding dorsal or ventral outlines of head; facets individually convex. Ocelli large, ocellar tubercles low. Ventral outline of head continuous, very slightly concave, only apex (fused bucculae) and basis (association with neck) outstanding. Dorsal ocular index 6.0, ventral ocular index 4.0. Antennae moderately long, thin; first segment strikingly long, cylindrical; segments 2-4 terete (not flagelliform, neither segment 4 subfusiform); antennal formula (longest segment first) 2, 3, 4, 1; segment 2 being 1.11 times as long as segment 3. Labium (Fig. 2) moderately long, rather thin, directed anterad (segments 1, 2) and ventrad (segments 3, 4), without particular structures, labial formula (longest segment first) 3, 2, 4, 1. Segment 3: 2.2 times as long as segment 2. Ventral outline of segment 2 emarginate near base and at midlength. Labrum reaching to middle of segment 2. Pronotum (Figs. 1-2). Collum short, 2.45 times as wide as long, with narrow precollum, dorsum with a linear impressed median area and pair of low and broad elevations, lateral area with low ‘pleural’ tubercle not visible in dorsal view. Collar constriction sharply delimited. Midlobe (dorsal side) with a linear, nearly percurrent median impression terminating just in front of posterior margin; disc with markedly plastic relief, with i) an inversely triangular anteromedial depression, ii) broad, not distinctly delimited, subcircular posteromedial depression, and iii) paired deep lateral pits emitting a lateral depression each; lateral margins broadly convex, interrupted, slightly notched because of lateral pits; posterior margin entire, trisinuate, sublateral shallowly concave parts slightly depressed, medial convex part broadly rounded, without edge. No traces of Y-shaped impressions. Constriction between midlobe and hindlobe broad, sharply demarcated. Midlobe 2.5 times as long as collum, 1.2 and 1.45 times as long as hindlobe maximum and median length, respectively. Midlobe 2.0 times as long as wide. Hindlobe ample, its median twice as long as collum, indicated neither by ridge nor impression, lateral margins rounded, posterolateral angles (in strictly dorsal view) subrectangular; posterior margin bisinuate (tetrasinuate, if moderately protruding posterolateral angles are counted), medially broadly and shallowly concave, moderately convex sublaterally. Hindlobe 3.2 times as wide as medially long. ‘ Proepimeral lobe ’ (see ŠTYS & BAŇAŘ 2006) extensive, distinctly exceeding posterior prosupracoxale posteroventrad, but not enclosing fore acetabula. Mesoscutellum. Concave central part equilaterally triangular, produced in long, apically rounded mucro and included into a larger triangle due to lateral association with forewing grooves. Prothoracic coxa, prosupracoxale, proacetabula, and prosternum. For details see Figs. 4 and 6. Cx 1 situated within proacetabulum, the latter open anteriorly and closed elsewhere. Proximal parts of lateral, anterior, and most of mesal faces of Cx 1 fully enclosed and externally delimited by prosupracoxalia. Anterior prosupracoxale excessively developed, anteromesally extended, embracing Cx 1 anterolaterally, anteriorly, and anteromesally. Transverse anterior part of anterior prosupracoxale fused with ventral part of collum (forming a not fully understood system of anteroventral prothoracic evaginations); anteromesal part of anterior prosupracoxale directed mesocaudad, fusing with triangular probasisternum, and taking part in formation of its strengthened, horizontal, sharp-edged lateral sides, here termed ‘ prosternal strigilatory edge(s) ’ (see Discussion). This composite triangular probasisternum horizontal, clearly delimited by its sharp edges; prosternellum adhering to probasisternum, sinuate in sagittal plane, more dorsal than probasisternum, with only the posterior, rounded, tongue-shaped part posteriad to coxae visible in ventral view. Legs rather short. Fore leg (Figs. 3, 6-16) extremely stout, femur and tibia incrassate. Cx 1 (Fig. 8) conical, anteroventral face with dense, prominent, drop-like cuticular thorns (Fig. 9) resembling and arranged like rasping files. Antero- and especially posterodorsal proximal parts of fore coxa with dense rows of different cuticular processes (Fig. 10), resembling serrate rasping files, each of those with several apical teeth, thus very similar to Pseudohenschiella hauseri Baňař & Štys 2006 from Madagascar (cf. BAŇAŘ & ŠTYS 2006). Proximal region of Tr 1 very narrow, more so than robust distal part, both regions separated by concave impression; entire adcoxal (dorsal) surface deeply concave (accommodating distal part of Cx 1 during flexion of Tr 1 ) and delimited by sharp, lateral, free anterodorsal and posterodorsal edges (about as long as one third of tr-f junction). F 1 2.5 times as long as wide, with basidorsal angle forming subrectangularly produced, apically rounded dorsal extension, the latter rising strikingly above dorsal surface of Tr 1 , and as high as its proximal diameter. Knee. Distinct remnant of a tripartite neopatella * (new term) visible dorsally in intersegmental F 1 -Ti 1 membrane when tibia maximally bent towards femur (Fig. 7). Ti 1 broadly triangular, 2.1 times long as wide, compressed in anteroposterior plane, both anterior and posterior faces each with vaguely delimited longitudinal depression. Cleaning comb short, formed by tightly packed short setae; three ventralmost spiniform setae longer and stouter. Intersegmental tibiotarsal membrane forming an evaginated pocket stretching far ventrad beneath tarsus itself (enabling apparently its close appression towards distal margin of tibia). Distiventral, armature-bearing process absent. Apicitibial armature (Fig. 15) consisting of seven spiniform setae: two short ventral (straight), three long subventral (all slightly oblique towards tarsus), and two short dorsal (strongly oblique towards tarsus). Ta 1 cylindrical, 1.7 times as long as wide, ventral surface slightly concave, tarsal armature (Fig. 16) of 1+1 proximal, curved spiniform setae and 1+1 distal setae (anterior one semicircular, posterior one broadly spiniform, shorter than proximal setae). Claws all of same shape, regularly curved, posterior one shorter and narrower. Fore leg sensilla on coxa, femur, and trochanter. Basal rim of Cx 1 anteromesally with coxal rim organ (Fig. 8), consisting of cluster of several (5-7) differently directed, straight setae and one distant short seta. Condylar trochanteral organ (Fig. 8) consisting of several (six?) poorly visible short setae. Anterior trochanteral organ (Figs. 8, 11) consisting of 6+1 campaniform sensilla (group of six, one isolated; posterior trochanteral organ (Figs. 13-14) consisting of six campaniform sensilla (five in straight row, one isolated). Anterior femoral organ (Figs. 8, 12) consisting of 3+1 campaniform sensilla (group of three and one isolated) very close to base of F 1 . Neopatella (new term) = sclerite(s) situated within the dorsal section of the F-Ti intersegmental membrane in a position where the original patellar limb segment or its remnants would be situated (see Discussion). In fore leg of female S. loebli sp. nov., the dorsal apex of F 1 is concavely excised and the small dorsolateral projections of its posterior margin provide articulation with the neopatellar articulatory tubercles.The area between these projections is filled with a tripartite neopatella, its medial part forming a narrow arcuate strip adhering to the concavity of the apex of F 1 , and its lateral parts being formed by strongly sclerotized, prominent, dorsally projecting articulatory tubercles associated with the medial strip of neopatella and articular projections of F 1 . The distal margin of the neopatella is associated with a short intersegmental F-Ti membrane; the latter separates the neopatella from a produced and feebly delimited process of basidorsal margin of Ti 1. Middle leg and hind leg. Lateral (dorsal, adfemoral) face of Cx 3 flat, largely covered by sandpaper-like file similar to that on Cx 1 . F 3 moderately incrassate, its basidorsal margin arcuate, clearly extending above the level of dorsal face of Tr 3. Proximal segments otherwise without particular structures, anterior and posterior faces of Ti 2 and Ti 3 sulcate. Ta 2 and Ta 3 two-segmented, segment 1 extremely short, without dorsal surface in lateral view (dorsal part of segment 1 visible as dorsal part of basitarsal ring filling up apex of tibia in anterodorsal view). Apices of both middle and hind tibiae each with strikingly short posteroventral and anteroventral setal combs; every comb terminating ventrally with a long spiniform seta. Claws and parempodial setae isomorphic. Forewing as usual for Systelloderes. Pterostigma strikingly well formed, long and wide, RP arising from its middle; rp-mp (= the anterior crossvein) entering open discal cell strikingly more distad than CuA3+4 (= the posterior ‘crossvein’); AP in claval area not developed. Ventral side of abdomen (distorted in the holotype) with series of 1+1+1 large sclerites on ventrites 3-8. Terminalia (distorted in the holotype). Posteromedial part of ventrite 7 thickened. Ventral laterotergite 8 distinct from subgenital plate, the latter strongly sclerotized, elongate, basal margin convex, distal margin concave in front of proctiger. Etymology. Dedicated to Ivan Löbl (Genève), an eminent coleopterist, our friend, and collector of the species. Bionomics. The holotype was collected by sieving leaf litter in a patch of a primary tropical New Caledonian rainforest surrounded by a secondary forest (I. Löbl, in epist.). Distribution. Known only from the type locality at Mount Koghi on the Grande Terre of New Caledonia.Published as part of Štys, Pavel & Baňař, Petr, 2007, The first species of Systelloderes (Hemiptera: Heteroptera: Enicocephalidae) from New Caledonia, pp. 3-15 in Acta Entomologica Musei Nationalis Pragae 47 on pages 4-11, DOI: 10.5281/zenodo.450356

Comment On The Proposed Precedence Of Bolboceras Kirby, 1819 (July)

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