PROC. ENTOMOL. SOC. WASH. 110(4), 2008, pp. 1220–1232

A NEW SPECIES OF TORNOCRUSUS (: : AENICTOPECHEIDAE) FROM ECUADOR, AND MORPHOLOGICAL NOTES

PAVEL Sˇ TYS AND PETR BANˇ ARˇ

(PSˇ) Charles University in Prague, Faculty of Science, Department of Zoology, Vinicˇna´ 7, CZ-128 44 Praha 2, Czech Republic (e-mail: [email protected]); (PB) Moravian Museum in Brno, Department of Entomology, Hviezdoslavova 29a, CZ- 627 00 Brno, Czech Republic (e-mail: [email protected])

Abstract.—Tornocrusus lattini Sˇ tys and Banˇarˇ n. sp. (Hemiptera: Heteroptera: Enicocephalomorpha: Aenictopecheidae: Aenictopecheinae) is described from Ecuador (-, U) and differentiated from other species of the genus. The species is characterized by several autapomorphies; the most important being the asymmetry of female first valvulae—the first somatic asymmetry found in the Enicocephalo- morpha. The forewings are characterized by presence of remigial CuA2 and claval AP, features here reported for the first time, but possibly overlooked in other Aenictopecheinae. Key Words: , forewing venation, foreleg architecture, male and female genitalia

The genus Tornocrusus Kritsky, 1977 topecheidae is a relic basal group of the (Enicocephalomorpha: Aenictopecheidae: true bugs and, therefore, any new struc- Aenictopecheinae; see Sˇtys 1995) was tural observations on it are of importance, established by Kritsky (1977) for an but that no comparative treatment is insufficiently described species, T. stysi attempted here. The new species described Kritsky 1977 from Costa Rica. The genus below differs in several important respects was revised by Wygodzinsky and Schmidt from Tornocrusus as characterized by (1991), who added, thoroughly described, Wygodzinsky and Schmidt (1991), but and illustrated six other new continental shares with the genus its general facies, Neotropical species, briefly mentioned basic features of forewing and hindwing one undescribed one, and transferred venation, basic features of male and Lomagostus aguilari Villiers 1978 from female terminalia, and several important Guadeloupe (Villiers 1978) into the genus. synapomorphies (pattern of foreleg arma- Since then, Sˇtys (2002) listed and keyed ture; inner (5 anterior) foretarsal claws the genus and considered it in a biogeo- extremely long, narrowing and filiform, graphical context (2008). In this paper, we whereas outer (5 posterior) foretarsal describe a new species of Tornocrusus claw abbreviated to reduced; mid- and from Ecuador, add observations to its hind tibiae with one apical comb each and generic diagnosis, list and compare all the one spinifiorm seta situated outside the known species, and add data on morphol- comb; propleural region reduced). There- ogy. It should be understood that Aenic- fore, prior to necessary comparative VOLUME 110, NUMBER 4 1221 morphological studies, we prefer to be Tornocrusus Kritsky, 1977 conservative, as we did in the recent case Tornocrusus Kritsky, 1977: 161. Type of a new Systelloderes Blanchard 1852 species: Tornocrusus stysi Kritsky, ˇ from New Caledonia (cf. Stys and Banˇarˇ 1977, by original designation. 2007), and retain the new species within Tornocrusus: Wygodzinsky and Schmidt, Tornocrusus. 1991: 51–85; Sˇtys 2002: 350, 362, 363. Dedication.—This paper, as well as the name of a new species of Aenicto- The following notes pertain to the pecheidae (Heteroptera: Enicocephalo- generic diagnosis (character states not morpha) hereby described, is dedicated mentioned by Wygodzinsky and Schmidt to Jack Lattin on the occasion of his 80th 1991, or found to be at variance). birthday, in recognition of his contribu- (1) The elevated interocellar part of the tions to the knowledge of Heteroptera, posterior lobe of the head in Torno- and, especially, for inspiring and attract- crusus spp. should be covered by a ing a number of students, many of which mat of ‘‘microtrichiae’’ (5 probably are nowadays eminent hemipterists. And short macrotrichia; PSˇ&JB), giving for being a good, long-standing friend— the area a dull appearance that Pavel. contrasts with the shiny surface of the rest of head (cf. Wygodzinsky MATERIALS AND METHODS and Schmidt 1991: 52, fig. 39A). Diagnostic characters.—The character This autapomorphic generic charac- states of the new species below described ter is missing in T. lattini sp.n. are compared with those of other Torno- (2) In the Enicocephalomorpha, the crusus species treated (excepting the longitudinal axis of the grasping often unavailable male or female) by foretibia coincides with that of the Wygodzinsky and Schmidt (1991). How- foretarsus, or the latter is adverging ever, we are not attempting a revision of to the distal edge of the foretibia. the genus, and, consequently, we are not However, the axes concerned are confident about which character states distinctly and strikingly divergent found to be at variance with Wygod- in T. lattini sp.n. and, consequently, zinsky and Schmidt (1991) are autapo- the long filiform claw of the fore- morphies of the new species and which tarsus is directed outwards (laterad) are based on overly generalization and instead of inwards (mesad). The incorrect or incomplete observations by tarsus cum posttarsus appears to Wygodzinsky and Schmidt (1991). have undergone torsion of nearly Forewing venation. Terminology ap- 180u. It is possibly an autapomorphy plied to forewing veins and cells is shown of T. lattini because the feature was in Fig. 11 and is based largely on that observed in all four specimens but used by Wygodzinsky and Schmidt was not mentioned or illustrated in (1991: fig. 6A) taken from Sˇtys (1980). any Tornocrusus species by Wygod- The names of cells and symbols used for zinsky and Schmidt (1991). the venation of clavus follow those of an (3) All FWs of T. lattini are provided unpublished study by J.Kukalova´-Peck with a longitudinal vein—called and P. Sˇtys, and are self-explanatory. provisionally CuA2 by us. It is Abbreviations.—FW—forewing; L— diverging from the stem M + Cu at length; l—left; max—maximum (maxi- about the level of apex of the claval mal); min—minimum (minimal); r— cell, runs parallel to the claval right; W—width. furrow and disappears proximad to 1222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

cu-aa. Wygodzinsky and Schmidt brown. Labium brown, segments 3 and (1991: figs 45F, 46C, 47H, 49N, 4 much paler apically; legs and abdomen 51H I, 53A, 55G) did not illustrate light brown. The external (posterior) face this feature in any Tornocrusus of forefemur and fore tibia of female species. No author studying the covered by greyish circular matt spots, aenictopecheids or enicocephalids some of them associated with a particu- has mentioned or illustrated such a lar macrotrichium, others without such vein (see Discussion). an association. Head and legs lustrous, rest of body Tornocrusus lattini Sˇ tys and Banˇarˇ, matt. Anterior lobe of head with small new species cuticular granules; posterior and dorsal faces of forefemur each with 1 row of Type locality.—Ecuador, Napo Prov- larger cuticular granules. Middle third of ince, NW of El Chaco, 1680–1690 m anterior face of foretibia, anteroventral a.s.l. Type material (locality labels cited part of forefemur and posterodorsal face exactly). Holotype: male—Ecuador, of forecoxa with numerous rows of Prov. Napo (21) 4.4 km NW of EL tilelike cuticular processes, resembling CHACO, S 00u189480 W77u509210 serrate rasping files. 28.xi.2006; 1680 m, YPT, M. Fika´cˇek & Vestiture yellowish. Setae on all faces J. Skuhrovec lgt. Cardmounted holotype of head semierect, curved anterad, longer to be deposited in the collection of on dorsal face of anterior lobe. Collum QCAZ Museum of the Pontifica Uni- covered with long, prominent setae on versidad Catolica del Ecuador, Quito, dorsal face and with shorter, semierect Ecuador. setae on lateral and ventral faces; all Paratypes: one male, same data as setae on collum directed anterad. Rest of holotype, one male and one female— pronotum and mesoscutellum with Ecuador, Prov. Napo (20) 6.5 km NW of prominent setae, directed caudad. Abdo- EL CHACO, S 00u189310 W77u509420 men regularly covered with semierect 28.xi.2006; 1680 m, sifting, M. Fika´cˇek setae, without long macrotrichia, with & J. Skuhrovec lgt., deposited in collec- exception of terminalia (see below; tions of Pavel Sˇtys and Petr Banˇarˇ, Figs. 19, 20). Antenna: all segments with preserved in alcohol. semierect, dense setae, distal part of Etymology.—lattini, a patronym de- segment 2 with several (6–8) longer, rived from the surname of Prof. John outstanding setae, as long as segment (Jack) D. Lattin (Corvallis, Oregon), an diameter; segments 3 and 4 with long eminent American hemipterist. (segment 3 with 12–15, segment 4 with ca Habitat and collecting methods.— 20) long, fine outstanding setae about Two specimens of T. lattini were sifted twice as long as segment diameter. from forest litter, and two specimens Labium with dense, semierect pilosity, were caught in a yellow pan trap, the first longer on dorsal face. Foreleg with 2 published example for enicocephalomor- types of setae: (a) long, prominent phans. trichobothria-like setae (tr-setae); (b) Description.—Measurements. Total shorter, semierect, denser setae. Vestiture length from 3.45 in male to 3.75 in female of foreleg markedly more dense in males. (other measurements see Table 1). Coxa and trochanter antero- and postero- Coloration and sculpture. Head, pro- ventrally with numerous semierect setae, notum, scutellum and FW dark brown. coxa with 4–5 tr-setae; trochanter with Antennal segments 1 and 2 brown, 6–8 tr-setae. Femur and tibia covered antennal segments 3 and 4 yellowish with semierect setae on all faces, more VOLUME 110, NUMBER 4 1223

Table 1. Measurements of Tornocrusus lattini (in mm).

Holotype—male Paratype—male Paratype—female Length antennal segment I 0.155 0.166 0.222 Length antennal segment II 0.277 0.290 0.366 Length antennal segment III 0.311 0.311 0.355 Length antennal segment IV 0.266 0.277 0.355 Total length antenna 1.009 1.044 1.298 Length head 0.422 0.444 0.577 Width head across eyes 0.355 0.377 0.466 Length eye 0.133 0.133 0.133 Width eye 0.082 0.084 0.077 Distance eye-apex of antennifer 0.044 0.044 0.088 Dorsal synthlipsis 0.191 0.200 0.288 Ventral synthlipsis 0.111 0.111 0.266 Interocellar distance 0.066 0.066 0.111 Distance ocellus—lateral margin of 0.077 0.077 0.088 posterior lobe head Width hind lobe head 0.344 0.355 0.444 Length hind lobe head 0.200 0.222 0.266 Length labial segment I 0.066 0.066 0.055 Length labial segment II 0.088 0.092 0.077 Length labial segment III 0.166 0.180 0.190 Length labial segment IV 0.133 0.144 0.177 Total length labium 0.453 0.482 0.499 Maximum length pronotum 0.333 0.377 0.477 Width pronotum mid lobe 0.466 0.488 0.555 Width pronotum hind lobe 0.533 0.577 0.666 Length fore femur 0.600 0.600 0.777 Width fore femur 0.244 0.266 0.377 Length fore tibia 0.511 0.580 0.688 Width fore tibia 0.155 0.166 0.233 Length foretarsus 0.190 0.200 0.200 Width foretarsus 0.080 0.080 0.133 Length anterior foreclaw 0.266 0.266 0.280 Length hind femur 0.844 0.888 0.866 Width hind femur 0.114 0.116 0.222 Length hind tibia 0.988 1.022 1.155 Width hind tibia 0.055 0.066 0.111 Total length 3.466 3.566 3.733 densely on posterior faces. Femur of male and 8–10 tr-setae; tarsus of female sparse- on dorsal face with 8–10 tr-setae, on ly covered with 2–4 tr-setae only. Mid- ventral face with14–16 tr-setae; femur of and hindlegs densely covered with semi- female on dorsal face without tr-setae, on erect setae on all faces, and longer, ventral face with 5–7 tr-setae. Tibia conspicuous setae, especially on postero- completely covered by semierect setae, dorsal faces. with exceptions of smooth apical third of Vestiture of FW. No microtrichia; anterior face and proximal third of occurrence of macrotrichia limited to ventral face; dorsal face in male with 8– veins; the distributional pattern of the 10 tr-setae, female without tr-setae, ven- latter, as follows. Marginal vein with two tral face in male with numerous (. 20) tr- (proximal sector) to irregularly three (up setae, in female with 8–10 tr-setae. Tarsus to the end of pterostigma) rows, the of male densely covered by semierect setae marginal ones long, the submarginal 1224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1–4. Tornocrusus lattini (photographed in alcohol). 1, 2, Male paratype. 1—General facies; total length 3.55 mm; note the position of the foretarsus against the foretibia. 2—Pronotum and pterothorax (partim). 3, 4, Female paratype. 3, General facies; length 3.73 mm; note the position of the foretarsus against the foretibia. 4, Pronotum and pterothorax. ones and those on the pterostigma very veins and to proximal sector of CuA2 as short; the ambient vein with one row; the well. Claval veins: AA1 + 2, AA3 + 4 and setae from the termination of ptero- AA with two rows of short setae, AP stigma up to the apex of wing very long, with 3–4 long, black setae. HW bare. moderately curved, then (following the Structure: Head (Figs. 1, 3) longer than vein proximad) the setae becoming grad- pronotum, ratio 1.18–1.27 in males, 1.20 ually shorter to minute (when the vein in female. Eyes slightly smaller in female, become submarginal and merges with Cu ratio eye L: distance eye to apex of + AA). The discal remigial veins each antennifer 3.02 in males, 1.51 in female. with one row of usually short setae (long Ratio max dorsal W eye : dorsal synth- on proximal sector of R only), becoming lipsis 0.42–0.50 in males, 0.27 in female, scarce to absent towards the FW apex; ratio max ventral W eye : ventral synth- presence of macrotrichia applies to cross- lipsis 1.1–1.2 in males, 0.38 in female. VOLUME 110, NUMBER 4 1225

Anterior and posterior lobes of head concave, especially proximal half. Neo- separated in males by conspicuous groove, patella absent. Foretibia three-dimen- in female groove indistinct. Ratio L to W sional apically, with an antero-dorsally of hind lobe of head 1.60–1.72 in males, directed ridge along apical third. Fore- 1.67 in female. Ocelli separated by about tibial bristle comb consisting of 33–34 1.5 times their width in males, 2.5 times setae in males and 33 setae in female. their width in female. Ratio interocellar Foretibial armature (Figs. 7–8, 22) con- distance to distance ocellus-lateral margin sisting of two groups of spines: a) of posterior lobe of head 0.86 in males, anterior one of two stout, semicircular 1.26 in female. Antenna longer than head spines and one conical spine; b) posterior and pronotum combined, 1.27–1.34 in one of three conical spines of unequal males, 1.23 in female, antennal formula length, the first and third markedly (longest segment first): iii-ii-iv-i in males, directed towards tarsus. The isolated iii-ii 5 iv,i in female. Segments i and ii seta on protruding tubercle, located wider than segments iii and iv, segment ii behind the dorsalmost seta of bristle widening distad. Ratio L of antennal comb occurring in both sexes. Foretarsus segment iii to L of segment ii 1.07–1.13 with four spines (Figs. 7, 9, 23), the two in males, 0.97 in female, ratio L of anterior of normal size, stout (proximal antennal segment iii to segment iv 1.12– one conical, distal one half-moon 1.17 in males, 1.00 in female. Labium shaped), two posterior weak, slender. stout, labial formula (longest segment Foretarsus with two claws, anterior one first): iii-iv-ii-i. long, irregularly filiform, posterior one Pronotum (Figs. 2, 4) weakly three- short and stout, about one third as long lobed, lateral margins diverging caudad, as anterior one. Unguitractor plate more markedly in male than in female. strikingly asymmetrical. Foretibal arma- Collum relatively well demarcated poste- ture, foretarsal armature, and shape of riorly by 1 + 1 sulci. Hindlobe with lateral foreclaw as typical for the genus. The margins nearly parallel (proximal two- long axis of the foretarsus diverging from thirds), with shallow median longitudinal that of the foretiba ectad, also the longer depression, defined by sulci. Hindlobe claw directed ectad. short, indistinctly demarcated from mid- Mid- and hindlegs stouter in female lobe. Ratio pronotum max L to max W than in males, ratio L to W of hindfemur 0.62–0.65 in males, 0.72 in female. Ratio 7.40–7.65 in males, 3.90 in female. Apex W of hind to W of midlobe of pronotum of mid- and hindtibiae (Fig. 10) with one 1.14–1.18 in males, 1.2 in female. Mesos- arc-shaped bristle comb and one large cutellum with sinuous posterolateral apical spine, outside the comb, in the edges, in dry-mounted specimens with middle of the ventralmost part of tibia in diagonal wrinkles (deformation after both sexes. mounting on cards). Propleuron striking- Forewing (Figs. 1, 3, 11, 14–17). Mac- ly reduced (cf. Wygodzinsky and Schmidt ropterous. Costal fracture squared; the 1991), ‘‘proepimeral lobes’’ minute, not crossvein r-m veinlike (nearly as long as reaching behind proacetabula. m-cu), hence, apex of axial cell truncate; Foreleg (Figs. 1, 3, 5–6) stout, ratio L basal cell strikingly short and narrow; m- to W of forefemur 2.26–2.46 in males, cu situated distinctly more basad than cu- 2.06 in female, ratio L to W of foretibia an. M + CuA branching in CuA1 and 3.30–3.50 in males, 2.95 in female. CuA2, the latter not reaching cu-aa. Short Forecoxa very long, about two thirds of AP distinct at the basis of AA3 + 4. foretibial length, trochanter without spe- Additional observations on FW. The cial features. Anterior face of forefemur major features and diagnostic character 1226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 5–10. Tornocrusus lattini. 5, 7–9, Male paratype. 6, 10, Female paratype. 5, Right foreleg, posterior view; note the filiform longer claw. 6, Right foreleg, anterior view, the filiform part of the longer claw broken off; note the dark circular spots on femur and tibia. 7, Right foreleg, tarsus and apex of tibia, antero-lateral view. 8, Right foreleg, tibial armature, posterior view. 9, Right foreleg, tarsal armature, and strongly asymmetrical unguitractor plate, antero-ventral view. 10, Apex of right hindtibia, ventral view; note the location of the spiniform seta outside of and distad to the comb. VOLUME 110, NUMBER 4 1227

Figs. 11–17. Tornocrusus spp. 11, Tornocrusus incaicus (modified from Wygodzinsky and Schmidt 1991, Fig. 46C, by addition of terminology)—model of FW venation of Tornocrusus spp. as known so far (i.e., lacking CuA2 and AP—compare Fig. 17). 12–17, Tornocrusus lattini. Schematic, without scale. 12, 13, Female paratype; costal fracture of the right forewing, stretched (12) and bent (13). 14–16, Aberrant forewing venation of male paratypes (CuA2 not shown). 14, Left FW; cubital cell once subdivided. 15, Right FW; cubital cell twice subdivided. 16, Left FW of another paratype; stigmal cell once subdivided, r- m plate-like, apex of axial cell pointed. 17, Male paratype, scheme showing the position of CuA2 and AP (the latter shown also as inset)—compare with Fig. 11. states as shown by Wygodzinsky and branching in about its midlength (distad Schmidt (1991: e.g., fig. 53A for T. stysi). to apex of anal loop) in M + CuA1 and Anterior margin slightly indented at the an independent vein CuA2 running large, square-shaped costal fracture. The parallel anterad to clavus, losing its ‘‘anchor’’ (5 a crossvein-like structure of veinlike character and diffusing into lower surface connecting basis of R with vaguely delimited sclerotization at about M + Cu stem) strikingly long. Preradial the level of claval organ, the sclerotized furrow percurrent along the submarginal area nearly reaching cu-aa (the latter cell. Pterostigma conspicuous, narrow, connecting Cu1a and AA). Anal cell elongate. The basal stem M + Cu (formed by AA1 + 2 and AA3 + 4) very 1228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 18–24. Tornocrusus lattini 18–20, 22–24, Male paratype. 21, Female paratype. 18, Phallus, and basis of r paramere (tilted to right), ventral view; lettering: ba—basal phallic apodeme; rpm—right paramere. 19, Terminalia (ventrite viii, pygophore, paramere, phallus), subventral view; elements of the right side and short microtrichiae omitted; parameres slightly spread (embracing and nearly covering the phallus when at rest); length of segment viii 0.24 mm, maximum length of pygophore 0.21 mm, maximum width of pygophore 0.22 mm. 20, Terminalia (pygophore, proctiger /5 abdominal segments x and xi/, parameres), dorsal view, short microtrichia omitted, parameres spread; the semisclerotized posteromedial region of dorsum ix is actually proximal part of dorsum x. 21, First valvifers, ventral view. 22, Right foretibia, apicitibial armature, posterodorsal view. 23, Right foretarsus, tarsal armature, anterior view. 24, Right foretarsus, tarsal armature, anterior view. short; AP recognizable as a short basi- lines in proximal stem and M + Cu marginal sector adjoining AA3 + 4. (distad to divergence of CuA2), a single, ‘‘Black lines’’ of FW. Some FW veins posterior line in proximal part of CuP; are characterized by presence of conspic- claval veins with a very conspicuous line uous, sharply delimited, straight and along anterior margin of AA1 + 2 and linear black lines along their margins joint basal arch of AA. Fluctuating (see, e.g., the inset to Fig. 17). Distribu- variation in FW venation (Figs. 14–16). tion: Marginal vein with two lines, Male paratype I: l FW (Fig. 16): marginal and submarginal, up to the stigmal cell subdivided into two equally start of preradial furrow; R with a single, long cells by a long, arcuate crossvein proximal, posterior line; M + Cu two running anterodistad; rp-m plate-shaped VOLUME 110, NUMBER 4 1229

(not veinlike), hence apex of axial cell senting a vaguely delimited basis of pointed. Male paratype II: l FW segment x fused with the membrane of (Fig. 14): CuA1b, before reaching the the original dorsum of the pygophore. ambient vein, branching in two sectors, Proctiger (segments x, xi) long, form- each reaching the ambient vein separate- ing a conspicuous tube, in untreated ly. r FW (Fig. 15): the same, and another specimens diagonally raised at an angle cross-vein running from the branching of about 35%, reaching the level of the point of CuA1 in CuA1a and CuA1b to apices of terminalia. Segment x of three the ambient vein—the cubital marginal sectors: (a) horizontal (see above), plate- cell thus subdivided in three cells. shaped, vaguely delimited, (b) and (c) Female paratype: Pterostigma less dis- raised, tubular, only slightly narrowing tinct, its posterior margin diffuse on both distad; (b) short, fully sclerotized, delim- l and r FWs (sexual dimorphism?). ited from (c) by a ring-like, circular Hindwings (Figs. 1, 3) of a modal apodeme; (c) ventral and lateral part type, exactly as illustrated by Wygod- forming a trough, dorsal surface mem- zinsky and Schmidt (1991: fig. 49F) for branous. Segment xi membranous, with T. pennai, only Cu and AA more distinct. 1 + 1 narrowly lunulate sclerites in lateral Also hamus (5 anterior, inner process / walls. 5 reduced vein/ of the ventral cell- Phallus inversely T-shaped, its basal delimiting vein) identically developed, transverse apodeme simple, fused with as a short recurrent hook contacting the posteroventral margin of the pygo- intracellular furrow. Pilosity lacking. phore, nearly as long as this margin and Abdomen (Figs. 1, 3) well sclerotized, about as long as the fully and perma- more weakly in female than in males, nently protruding body of the phallus without patches of specialized structures. itself. The body tubular, moderately Male terminalia (based on male narrowing distad, strongly sclerotized, paratype) (Figs. 18–20): Vestiture in- delimited from the transverse basal cluding some extremely long, thin, out- apodeme by a thick, C-shaped, ventral standing macrotrichia: 1 + 1 on ventrite semi-ring, and subdivided in three parts: viii, 4 + 4 on ventral and lateral sides of (a) proximal, fully sclerotized tube; (b) the pygophore (these somatic macrotri- mesal, equally long trough, with ventral chia about as long as segment diame- surface membranous; apically pointed, ters); about 4–5 on each paramere not bifid, (c) apical, slightly distended, (exceeding the paramere diameter). membranous ‘‘glans’’ provided with Segment viii completely free, its ter- large ventral aperture. The sectors (a) gum and ventrite equally long, subquad- and (b) penetrated by a duct directed rate, without particulars. Pygophore towards the aperture and provided with (segment ix) free, not at all telescoped finely serrate margins. No phallic pro- within segment viii, trough-shaped, its jections (cf. Wygodzinsky and Schmidt anteroventral margin medially emargin- 1991) visible in lateral view. ate, the ventral sector convexly vaulted, Parameres large and broad, in ventral tergum (5 dorsal bridge of the pygo- view (untreated specimens) overlapping phore) absent, its area membranous, and covering the phallus, directly asso- lacking macrotrichia. The sclerotized ciated by their desclerotized ventral bases dorsolateral parts of the pygophore with lateral parts of the basal apodeme, covering (dorsal view) about one fifth without any intervening connective of its dorsal width, their mesal margins structures but mobile owing to their irregularly dentate, asymmetrical. The ventral desclerotized bases. Parameres dorsal posteromedial area pilose, repre- symmetrical, irregularly racquet-shaped, 1230 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with rounded outlines and apices, me- Furthermore, T. lattini differs from sally hollow to allow for a full embrace- nearly sympatric T. penai by smaller ment of the phallus; their apicidorsal size (male 3.45 mm 3 4.4–4.7 mm, surface provided with ‘‘two-segmented’’ female 3.75 mm 3 5.1 mm), absence of finger-shaped posteromesally directed dorsal projection on phallus, and sim- process allowing for locking the phallic pler shape of parameres. tube mesodorsally. Outline of a para- The small size, general shapes of mere strongly dependent on the angle-of- parameres and phallus, dimensions of view, in strictly lateral view seemingly pronotum, size and mutual distance of very narrow. ocelli, ratio width of eyes to width of Female terminalia (Fig. 21) (Not ex- dorsal synthlipsis and number and shape amined in detail): Ventrite vii not form- of setae in fore apicitibial comb combined ing a subgenital plate. Ventral part of differentiate the new species from every segment viii formed by two large, prox- other described American species (cf. imally partly fused first valvifers. First Wygodzinsky and Schmidt 1991). valvulae distinct, arising from the distal Macroptery is a good distinguishing parts of the mesal valviferal margin, character of all Tornocrusus species from proximally mutually fused, asymmetri- the only insular species, brachypterous T. cal: the right one terminating in a aguilari (Villiers 1978) from Guadeloupe. strongly sclerotized, sharp, mesally di- rected hook, the right one with a blunt, LIST AND DISTRIBUTION OF strongly sclerotized, terminal, rugulose TORNOCRUSUS SPECIES apex and additional minute sclerites Tornocrusus aguilari (Villiers 1978) (as (Fig. 21). Second valvifers (or ventral U laterotergites ix?) large, forming a distal Lomagostus), : 727, fig. 44; Wygod- pair of ventral plates, second valvulae zinsky and Schmidt, 1991: 65–71, fig. fully covered (but not embraced) by first 44; Lesser Antilles: Guadeloupe Is. valvulae. Tornocrusus browni Wygodzinsky and Proctiger strikingly different from that Schmidt 1991, U: 65, 71–73; figs. 38– of male: segment x, short, inconspicuous, 45; Brazil: Estado Sao Paulo. not at all raised, ball-shaped, formed by Tornocrusus incaicus Wygodzinsky and a pair of lateral sclerites, and dorsal and Schmidt 1991, -: 63,73, figs. 38, 46; ventral membranous parts, segment xi Peru: Cuzco minute, situated within the distal aper- Tornocrusus lattini Sˇtys and Banˇarˇ, sp.n., ture of x. -, U: Ecuador, Napo Province Differential diagnosis.—In qualitative Tornocrusus occidentalis Wygodzinsky character states, T. lattini sp.n. differs and Schmidt 1991, -, U: 63, 64, 73– from other species (or their known 77, Figs. 38, 47, 48; Colombia: Valle sexes) of Tornocrusus by the absence of Tornocrusus penai Wygodzinsky and a matt, pilose interocellar area (present Schmidt 1991, -, U: 63–64, 77–79, in all other species), foretarsus and figs. 38–42, 49–50; Ecuador: Napo- foreclaws diverging ectad from the apex Pastaza of foretibia (instead of converging to it Tornocrusus sp.n. cf. penai (undescribed): as in other species), segmentation and Wygodzinsky and Schmidt, 1991, -: unique shape of the apex of phallus, 78, fig. 50: Ecuador unique shapes of the apices of the first Tornocrusus sturmi Wygodzinsky and female valvulae and their asymmetry, Schmidt 1991, -, U: 63, 64, 78–83, and grey circular spots covering poste- figs. 38, 43, 51, 52; Colombia: Narono rior face of forefemur and foretibia. and Cundinamarca VOLUME 110, NUMBER 4 1231

Tornocrusus stysi Kritsky 1977: 161–168, remigial vein we call provisionally figs. 1–12; Wygodzinsky and Schmidt, CuA2 (Fig. 17) are certainly plesio- 1991: 63, 83, figs. 38, 53, 54; Costa morphic characters in Heteroptera, Rica and we strongly doubt that they Tornocrusus venezolanus Wygodzinsky would occur solely in T. lattini.Itis and Schmidt, 1991, U: 65, 83–85, figs. more probable that the AP has been 38, 55; Venezuela: Estado Aragua overlooked in other macropterous Aenictopecheidae and CuA2 has not DISCUSSION been examined carefully being a priori relegated to a pigmented Some unusual features observed in adclaval strip. A reappraisal of Tornocrusus lattini and generalities re- venation in all Aenictopecheidae is sulting from its study are briefly dis- necessary but is beyond the scope of cussed below. this paper. Forewings: (a) The extensive costal fracture and Male and female terminalia: Shape briefly interrupted sclerotized costal and architecture of both are undoubtedly margin of T. lattini (Figs. 12–13) of great diagnostic value, which cannot allow for a sharp bending of FW be yet fully exploited because the genita- anterior margin as well as for its lia of both sexes are known only in T. torsion and nearly a flexion (distal lattini and T. penai. The general shape of part of FW turned over proximal parameres is probably species-specific part) starting at this pivoting point. (Wygodzinsky and Schmidt 1991) but it This function is apparent in alcohol- is difficult to describe it exactly since the preserved specimens, and probably parameres are fused with the phallic has been overlooked in those taxa of apodeme (the latter being in turn fused Enicocephalomorpha provided with with the posterior margin of pygophore), a costal fracture. cannot be dissected well in minute and (b) The great fluctuating variation in often unique specimens, and their ob- FW venation (Figs. 14–16) is char- served shape is obviously strongly de- acteristic of all the Enicocephalo- pendent on the angle of observation. We morpha but is particularly pro- are not sure whether the shape of the nounced in the Maoristolinae (Sˇtys, phallus apex has the diagnostic value unpublished) and Aenictopecheinae ascribed to it by Wygodzinsky and (Sˇtys 1980; present study; Wygod- Schmidt (1991) because it probably zinsky and Schmidt 1991). Such a depends on the functional state of the variation seems to be characteristic organ. of the basal clades of true bugs, such On the other hand, the shape of the as the Enicocephalomorpha and elements of female genitalia, particular- (Sˇtys, unpublished ly first valvifers, seems species-specific observations). Addition of veins is and easy to study. However, the anat- usually involved, although obligato- omy of female external genitalia must ry (?) losses of vein in individual be studied in greater detail (their unex- taxa are not infrequent in the pected complexity is suggested by Wy- Enicocephalinae, and have resulted godzinsky and Schmidt (1991: fig. 43F, in unsubstantiated erections of new photomicrographs of T. sturmi). We genera as discussed by Sˇtys (2002). stress that the asymmetry of the first (c) Presence of a rudimentary vein AP valvulae in T.lattini,asobservedinour on clavus and the most posterior female specimen and illustrated here 1232 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Fig. 21), is not only autapomorphic in Sˇtys, P. 1980. Australostolus monteithi, gen. n., sp. Tornocrusus species but also represents n.—First record of an Australian aenictope- a first case of somatic asymmetry in the cheine bug (Heteroptera: ). Acta Entomologica Bohemoslovaca 77: Enicocephalomorpha. 303–321. Forelegs: The grey spotting of female ———. 1995. Enicocephalomorpha, pp. 67–73. In forelegs and the torsion of the foretarsus Schuh, R. T. and J. A. Slater, eds. True Bugs against the foretibia with consequent of the World (Hemiptera: Heteroptera): Clas- direction of the claws outwards are sification and Natural History. Cornell Uni- versity Press, Ithaca and London. autapomorphies of T. lattini, but we are ———. 2002. Key to genus-group taxa of the not yet able to appreciate their anatom- extant Enicocephalomorpha of the world, their ical and functional significance. list, and taxonomic changes (Heteroptera). Acta Universitatis Carolinae Biologica ACKNOWLEDGMENTS 45(2001): 339–368. This project was financially supported ———. 2008. Zoogeography of Enicocephalomor- pha (Heteroptera). Bulletin of Insectology 61: by the Grant Agency of the Academy of 137–138. Sciences of Czech Republic No. IAA Sˇtys, P. and P. Banˇarˇ. 2007. The first species of 601110706 to the senior author. The Systelloderes (Hemiptera: Heteroptera: Enico- authors are obliged to Martin Fika´cˇek cephalidae) from New Caledonia. Acta En- (Prague) and Jirˇ´ı Skuhrovec (Prague) for tomologica Musei Nationalis Pragae 47: 3– collecting material of the new species in 15. Villiers, A. 1978. Sur quelques he´te´ropte`res de Ecuador, putting it at our disposal, and Guadaloupe (Henicocephalidae, ). providing field data. Annales de la Socie´te´ Entomologique de France (n.s.) 14: 727–734. LITERATURE CITED Wygodzinsky, P. W. and K. Schmidt. 1991. Kritsky, G. 1977. Two new genera of Enicocepha- Revision of the New World Enicocephalomor- lidae (Hemiptera). Entomological News 89: pha (Heteroptera). Bulletin of the American 65–73. Museum of Natural History 200: 1–265.