Research Article |
Corresponding author: Petr Baňař ( petrbanar@seznam.cz ) Academic editor: Alfred Wheeler
© 2018 Pavel Štys, Petr Baňař.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Štys P, Baňař P (2018) A new Xenicocephalus species from Ecuador (Heteroptera, Enicocephalomorpha, Enicocephalidae). In: Wheeler Jr AG (Ed.) A Festschrift Recognizing Thomas J. Henry for a Lifetime of Contributions to Heteropteran Systematics. ZooKeys 796: 33-47. https://doi.org/10.3897/zookeys.796.24538
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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.
Abdomen, biology, differential diagnoses, distribution, Ecuador, forelegs, Hemiptera , new species
A new Neotropical genus and species of Enicocephalidae (Enicocephalinae), Xenicocephalus giganticus Wygodzinsky & Schmidt, 1991, was well described and illustrated by
Owing to the specimen’s uniqueness, we could not study its anatomy in depth. Our descriptions of the head, antennae, pronotum and forelegs in an adult female, however, are the first for the genus. The female holotype of X. giganticus lacked these body parts, and the genus description had to be supplemented (
In the Discussion we emphasize some systematic issues concerning Xenicocephalus that have never been considered, comprehensively review certain aspects of biology, and briefly mention the construction of the fore leg of Xenicocephalus, which is unique among Enicocephalomorpha. Their architecture will be considered in a separate paper assessing the suprageneric phylogenetic classification of the Enicocephalomoropha (Štys and Baňař, in prep.).
The term ocular index refers to the ratio of the minimum interocular distance to the maximum width of the eye; it is best calculated if measured as twice minimum interocular distance/maximum width across the eyes, minus minimum interocular distance. Measurements were taken using a SZP 11 ZOOM stereoscopic microscope with an eyepiece micrometer.
Color photographs of the newly described species were taken with a Leica MSV266 camera. Scanning electron micrographs of a gold-coated left foreleg were taken using a JEOL 6380 LV scanning electron microscope.
Label data are cited verbatim, including potential errors, using a slash (/) to separate lines on the label; different labels are mentioned and indicated by a double slash (//). Our notes are in [square brackets].
For simplicity, our nomenclature for veins and cells follows that used by
Abbreviations used in the text:
DLTG dorsal laterotergite;
F female;
L larva of the fifth instar;
M male;
MTG mediotergite.
Xenicocephalus giganticus Wygodzinsky & Schmidt, 1991 by original designation.
Measurements (in mm) of female holotype; L = length; W = width. Total body L 8.60. Head: total L (without neck) 1.31; posterior lobe, L 0.51, posterior lobe, W 0.71; distance eye-apex of antennifer 0.37; maximum width across eyes 0.67; dorsal minimum interocular distance 0.33; ventral minimum interocular distance 0.20; eye L 0.22. Labium total L 0.60. Antenna: Segment I L 0.40; segment II L 0.96, segment III L 0.89, segment IV L 0.82. Pronotum: total L (maximum) 1.64; collum, L (median) 0.28, maximum W 0.78; midlobe, L (median) 0.64, midlobe, W (maximum) 1.42; hindlobe, L (maximum) 0.87, hindlobe, L (median) 0.40; hindlobe, W (maximum) 2.02. Foreleg: femur L 1.49, femur, maximum W 0.51, tibia L 1.36, tibia maximum W 0.29. Forewing L 5.95, W 2.18. Hindwing L 4.94, W1.92.
Coloration dark brown (Figure
Pilosity. Antennae densely covered with short, semi-erect setae, dorsal and lateral parts of head covered with long, semi-erect setae of variable orientation, mixed with shorter, erect setae similar to those on antennae and compound eyes. Venter of head with long semi-erect setae. Vestiture on pronotum, lateral, and ventral parts of thorax similar to that on dorsum of head. Forelegs with numerous long semi-erect setae of different orientation, setae on ventral face of foretibia shorter and erect. Forewing veins with short semi-erect to erect setae; hindwing veins bare. Dorsum of abdomen with semi-erect and erect setae, longest on “outer” laterotergites, becoming shorter to nearly absent toward medial parts of mediotergites. Lateral faces of laterotergites only with short, erect setae. Venter of abdomen densely covered with vestiture, becoming denser and longer towards apex of abdomen.
Texture. Body faces, including forewing, densely covered with countless cuticular microgranules, giving a matt appearance. Foretibia along almost entire ventral face (except proximal sixth) with two rows (posterior and anterior one) of irregularly placed large, semi-globular to slightly conical, strongly sclerotized granules (Figure
Head. Rather narrow, strongly elongate (Figs
Antennae. Insertion nearly subterminal, segment 1 with a large prescapite, segment itself strongly widening distad, much surpassing apex of head, segment 2 terete, slightly thickening distad, segments 3 and 4 long, thinner, but less than subflagelliform. Antennal formula (longest segment first): II:III:IV:I.
Labium. Short, reaching anterior margins of eyes. Labial formula (longest segment first) III-IV-II=I. The dorsal (morphologically ventral) outline of segment III moderately convex, ventral one straight.
Pronotum (Figure
Fore acetabula widely separated and widely open. Mesoscutellum broadly triangular, apex not produced but provided with transversely oval swelling with marginally radiating macrotrichia. Mesonotum and metanotum (Figure
Forelegs. Foretrochanter (Figs
Forefemur (Figs
Foretibia (Figs
Midlegs and hindlegs short and robust, first tarsomeres very short. Apices of middle and hind tibiae with two bristle combs, anterior and posterior ones each consisting of ca. 12–15 isomorphic setae, and two or three much longer, ventrally placed setae.
Post-tarsi could not be studied in detail. Fore post-tarsus with strikingly split unguitractor plate along median. Claws heteromorphic, strikingly slender and short, longest at foreleg (posterior claw reduced to stump), shortest at midleg. Fore claw and hind claws nearly straight, middle claw basally thick and distally regularly arcuate.
Forewing (Figure
Hindwing (Figure
Holotype female, labelled: 'Ecuador camino / Aloag-Tandapi / Pr. Pichincha 2600m / 12.II.1983 A. Roig [handwritten] // Drake Colln. ex / J. Maldonado C. / Coll. 1996 [printed] // Xenicocephalus / sp. nov. / P. Baňař det. 2014 [handwritten, partly printed] // HOLOTYPE / Xenicocephalus / tomhenryi sp. nov. / P. Štys & P. Baňař det. 2018 [printed red label]’. Dry-mounted, left foreleg and right wings mounted separately; right middletarsus missing. Deposited in Department of Entomology of National Museum of Natural History, Washington, D.C. (USNM).
Dedicated to our dear colleague Thomas J. Henry, eminent student of the Heteroptera, for long-standing cooperation and friendship. Pavel will always remember Tom´s and Katy´s hospitality and kind assistance during his stay at their house in Silver Spring, Maryland, after his mishap in 2014.
The following comparative paragraphs are intended to serve as a diagnosis and comparative diagnosis. Because of the paucity of material, we could not always determine whether the differences are species-specific, sex-specific, or represent individual variation. The last alternative may particularly involve characters of the forewing venation, which is notoriously variable and subject to teratological mutations (cf.
The data on Xenicocephalus species are organized as follows: (1) X. tomhenryi female from Ecuador (holotype), (2) X. josifovi male from Suriname (holotype), (3) X. giganticus female from Colombia (incomplete holotype), (4) X. sp., larva 5 from Colombia: (Santa Marta: San Sebastian de Marago) assumed by
Xenicocephalus tomhenryi sp. n., female holotype, left foreleg, scanning electron micrographs A coxa, trochanter and basis of femur, anterior view B coxa and trochanter, ventral view C femur, anterior view D tibia, anterior view E detail of ventral concavity of tibia F bristle comb of tibial apex, anterior view.
Antennae
(1) X. tomhenryi F – segment I long, strikingly thickening distad; II long, terete, slightly thickening distad, about as long as head.
(2) X. josifovi M – segment I short, thick, not thickening distad; II short, thicker but not thickening distad, about 1.5 times as long as head.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 – segment I short, thickening distad; II long, terete, not thickening distad, about as long as head.
Head (size of eyes sexually dimorphic?)
(1) X. tomhenryi F – preocular part of head long, eyes short, distances anterior margin of eye-insertion of antenna and posterior margin of eye-ocellus longer than maximum length of eye; postocular part of anterior lobe & transverse constriction strikingly long; ocelli submarginal.
(2) X. josifovi M – preocular part of head short, eyes long, distance anterior margin of eye-insertion of antenna and posterior margin of eye-ocellus much shorter than maximum length of eye; postocular part of anterior lobe not present, constriction narrow; ocelli marginal.
(3) X. giganticus F – ?
(4) X. giganticus (F?) L 5 – preocular part of head long, eyes minute, lateral, distance anterior margin of eye-insertion of antennae as long as maximum length of eye; postocular part of anterior lobe nearly as long as eye; constriction deep and narrow; ocelli not mentioned in original description, but their rudiments indicated (
Pronotum
(1) X. tomhenryi F – posterior margin of collum strongly convex, encroaching onto midlobe region; hind lobe “entire,“ median part not differentiated; anterolateral parts of hindlobe embracing posterolateral parts of midlobe, posteromedial notch of hind lobe ca. twice as deep as maximum median length of hindlobe.
(2) X. josifovi M – posterior margin of collum transverse; hindlobe “bipartite,” creating impression of two opposite leaves attached to broad and weakly sclerotized median region; anterolateral parts of hindlobe not extending cephalad, posteromedial notch of hind lobe ca. half as deep as maximum median length of hindlobe.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 – posterior margin of collum very moderately convex; mid- and hindlobes not differentiated.
Mesoscutellum
(1) X. tomhenryi F – broadly rounded, apex with transversely oval swelling with radiating marginal macrotrichia.
(2) X. josifovi M – amply triangular, apically mucronate.
(3) X. giganticus F – amply triangular, apically mucronate.
(4) X. giganticus (?) L 5 – 0.
Forewings (individual variation and most potential teratologies could not be assessed). We are not certain about the presence of AP in any Xenicocephalus species (contrary to our previous statement on X. josifovi (
(1) X. tomhenryi F – C&Sc, R and Rs extremely strongly thickened, anteradial furrow along edge of C⪼ veins delimiting base of discal cell (part of M and part of Cu proximal to cu-an) unequally long, M about three times as long as Cu; r-m vein-like; apex of discal cell close to wing margin, 2 short distal veins entering wing margin; fork Cu1a-Cu1b far distad to cu-an. Forewing macropterous, conspicuously exceeding apex of abdomen.
(2) X. josifovi M – C&Sc, R and Rs moderately thickened, anteradial furrow within subcostal cell; veins delimiting base of discal cell (part of M and part of Cu proximal to cu-an) equally long; r-m point-like; apex of discal cell close to wing margin,1 hardly distinct distal vein entering wing margin; fork Cu1a-Cu1b coinciding with position of cu-an. Forewing macropterous, exceeding apex of abdomen.
(3) X. giganticus F – C&Sc, R and Rs extremely strongly thickened but anteradial furrow within subcostal cell; veins delimiting base of discal cell (part of M and part of Cu proximal to cu-an) equally long; r-m vein-like; apex of discal cell distant from wing margin,1 distinct vein reaching wing margin; fork Cu1a-Cu1b far distad to cu-an. Forewing submacropterous, not exceeding apex of abdomen.
(4) X. giganticus (?) L 5 – 0.
Foretrochanter (perceived shape strikingly dependent on angle of observation).
(1) X. tomhenryi F – broadly rounded, apex with transversely oval swelling with radiating marginal macrotrichia.
(2) X. josifovi M – ventral side with prominent ridge terminating in small ventral tubercle exceeding base of femur ventrad and only inconspicuous distad.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 – ventral side with prominent strongly sclerotized ridge-like projection exceeding base of femur ventrad but not distad.
Forefemur
(1) X. tomhenryi F – strikingly thick and curved, arcuate, dorsal face convex, ventral face deeply and percurrently concave; groove delimited by two marginal rows of densely packed minute subglobular platelets of stronger sclerotization, anterior row of large granules more developed, on slightly elevated rim, granules in posterior row smaller, nearly subglobular proximally, becoming lens-like platelets distally, appearing in lower magnification as impression of two deeply brown to blackish lines. Surface of forefemoral concavity with countless lens-like platelets, heavily sclerotized, blackish, the surface of the concavity densely pilose. No other blackish platelets or granules present.
(2) X. josifovi M – distinctly curved, moderately C-shaped; ventral face concave, with vestiture lacking, parallel-sided and sharply delimited at anterior and posterior edges by row of macrotrichia and irregularly distributed black granules intermixed with row of conspicuous, high, non-setigerous conical tubercles.Ventral concavity with numerous small, broad, transverse scale-like structures. Blackish granules also on distal two thirds of lateral and dorsal faces.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 – conspicuously curved, bearing numerous cuticular granules dorsally and ventrally.
Foretibia
(1) X. tomhenryi F – thick, flattish, rather uniformly broad, very moderately arcuate, ventral face deeply percurrently concave and densely pilose, concavity about as long as that of forefemur, margins of concavity delimited as in forefemur by subglobular platelets. Foretibia along ventral face (except proximal sixth) with two rows (posterior and anterior) of irregularly placed large, semi-globular to slightly conical, strongly sclerotized granules, appearing in lower magnification as impression of two deeply brown to blackish lines. No other blackish platelets or granules present.
(2) X. josifovi M –cylindrical, of uniform width, only dorsal outline slightly curved. Entire ventral face moderately concave, vestiture lacking, edges of tibial concavity less sharply delimited than those of femoral one. Anterior edge with 14, posterior edge with numerous conical tubercles of same shape as on femur. Anterior face with ca. 50 black granules, posterior face with several hundred. Ventral concavity with numerous small, broad, transverse scale-like structures.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 –inner apical angle in form of pointed, strongly sclerotized projection bearing 3–5 straight, slender spines inserted below apex. Apex of central portion of inner surface of foretibia with field of short, stout setae.
Apicitibial and tarsal armature of foreleg
(1) X. tomhenryi F – tibial process long and narrow, strikingly differentiated from remainder of distal tibial edge, with four slender and more dorsal spines, and one ventral thick, short, conical spine from more ventral tubercle. Tarsal armature from four somewhat thicker setae (two anterior + two posterior = two dorsal + two ventral) distributed among normal macrotrichia, not longer than these, and recognizable only by presence of longitudinal grooving.
(2) X. josifovi M – tibial projection moderately large, rounded, with seven slender subapical spines, four in ventral row, three in dorsal row; tarsal armature from four spines, three of them very long and slender, one stout and conspicuously shorter.
(3) X. giganticus F – ?
(4) X. giganticus (?) L 5 – tibial projection acutangular but not markedly differentiated from distal edge of tibia; with 3–5 slender subapical spines.
Pregenital abdomen, dorsum (a comparative study required)
(1) X. tomhenryi F – complex lattice system (elevated rails and rungs)
present.
(2) X. josifovi M – lattice absent.
(3) X. giganticus F – lattice absent.
(4) X. giganticus (?) L 5 – lattice absent.
Nomenclatural and taxonomic notes.
We cannot be certain which interspecific differences (
The distribution of Xenicocephalus can be characterized as “southern continental Central America and northern South America” (
Only scant information is available on the biology of Xenicocephalus. For example, pteygopolymorphism (adults of the three species are submacropterous to macropterous) and swarming are unknown. The male X. josifovi from Suriname was taken at light, whereas the other specimens were collected accidentally or in pitfall traps in lowland to montane forest (to 2600 m). The scattered data on collection dates do not provide useful information. However, the peculiar and, among Enicocephalomorpha (and perhaps all other Heteroptera), unique shapes of the forefemur and foretibia in Xenicocephalus suggest a specialized and unique mode of catching and handling a certain kind of prey. We predict that the curved, raptorial forefemora and foretibiae, both provided with an extensive and deep concave area on their ventral face, are suited for holding and possibly cracking strongly sclerotized, convex and rounded prey (as in similarly shaped beetles).
The larvae of species of Xenicocephalus (cf.
This work was financially supported (to Petr Baňař) by the Ministry of Culture of the Czech Republic, as part of its long-term conceptual development program for research institutions (ref. MK000094862).