Nene, Kment, 2015

Kment, Petr, 2015, Two new genera of Madagascan Pentatominae (Hemiptera: Heteroptera: Pentatomidae), Acta Entomologica Musei Nationalis Pragae 55 (2), pp. 591-624 : 608-611

Nene gen. nov.

Type species. Nene undulatum sp. nov., here designated.

Description. Structure. Body ( Figs 6–8 ) wide, subquadrate in outline, robust and biconvex, more convex dorsally than ventrally.

Head ( Figs 44–45 , 48 ) large, slightly wider than long (1.0: 1.2), quadrate, directed forwards, its posterior part behind eyes encompassed by anterior pronotal margin ( Fig. 6 ). Vertex slightly convex, remaining surface of head nearly flat, sloping obliquely downwards, parallel to surface of anterior portion of pronotum ( Fig. 45 ). Compound eyes ( Fig. 44 ) very small, completely protruding from head outline. Postgenae (= tempora) behind eyes very narrow, narrowing laterad and not surpassing eyes laterally ( Fig. 44 ). Ocelli very small, situated posteromedially of eyes near anterior pronotal margin ( Fig. 44 ). Mandibular plates ( Fig. 44 ) very wide, explanate, slightly depressed in front of eyes, without anteocular spine or incision; straight in basal two thirds, very slightly diverging anteriad, rounded in apical third, meeting and surpassing each other in front of clypeus, leaving minute V-shaped incision anteriad of clypeal apex. Antenniferous tubercles small ( Figs 45 , 48 : at), situated immediately in front of eyes, concealed in dorsal view. Antenna pentamerous, slender. Antennal segments I, IIa and IIb cylindrical, segment I stout, segments IIa and IIb much more slender, segment III narrowly clavate, distinctly widening in its basal third, segment IV spindle-shaped. Bucculae ( Figs 45 , 48 : bc) low, bisinuate, arcuate and highest in anterior third, slightly concave in middle third, continually narrowing posteriad in wide arch in posterior third. Labrum ( Figs 45 , 48 : lr) flat and narrow. Labial segment I ( Figs 45 , 48 : lb1) slender, in lateral view completely covered by bucculae anteriorly, partly exposed posteriorly, slightly surpassing bucculae, not reaching posterior margin of head; labial segment III longest; apex of labium reaching base of ventrite IV.

Pronotum ( Figs 6, 8 , 46 ) transverse, its anterior margin concave to receive head. Anterolateral angles truncate, fitting posterior margin of compound eyes, each bearing a small denticle laterally ( Fig. 46 ). Anerolateral margins convex and rounded, carinate. Humeri ( Figs 6 , 46 ) large, very prominent laterally, flattened, subtriangular, apically nearly rectangular, slightly rounded, provided with a smaller, wave-shaped preapical lobe anteriorly (anterior margin gently arched, lateral margin perpendicularly truncate). Posterior margin of pronotum broadly arcuate from humeri to anterolateral angles of scutellum ( Fig. 46 ), nearly straight along base of scutellum. Disc of pronotum ( Fig. 8 ) horizontal posteriad of humeri, anteriorly obliquely sloping forward towards head, flat except for two very shallow depressions on base of humeri.

Scutellum ( Fig. 6 ) large, long and wide, nearly calyx-shaped; lateral margins of frenal portion slightly convex, narrowing towards apex, there sinuate; sides of postfrenal portion straight, slightly narrowing posteriad; outline of the apex in form of a wide pointed arch. Each anterolateral angle with narrow crevice-like C-shaped fovea with callose margins ( Fig. 6 ). Disc of scutellum ( Fig. 8 ) distinctly raised anteromedially, forming funnel-shaped gibbosity; its posterior part forming short longitudinal keel at level of apices of frena; apex flat.

Hemelytra ( Fig. 6 ). Clavus long and narrow, triangular, with about 4 irregular lines of punctures in its widest, basal part and only a single line of punctures in middle and distal parts. Costal margin of corium very shallowly concave in basal third, convex and regularly arcuately narrowing posteriad in its remaining length. Anterodistal angles of corium slightly surpassing apex of scutellum ( Fig. 6 ); both anterodistal and posterodistal angles widely rounded. Membrane short and wide, reaching (♀) or slightly surpassing (J) apex of abdomen; veins reticulate.

Thoracic pleuron and sternum ( Fig. 7 ). Meso- and metacoxae widely separated, distinctly more widely than procoxae. Prosternum narrow, groove-like, emarginated laterally by callose inner margins of procoxal cavity. Mesosternum with median longitudinal carina, anteriorly in form of wide and low triangle, narrowing posteriad and forming low, narrow, but well emarginated keel in middle of mesosternum, gradually disappearing posteriad. Metasternum flat. Surface of pleura ( Fig. 49 ) uneven, propleuron with one depression on base of both anterolateral and humeral angles, mesopleuron with two small depressions anterolaterally, metapleuron with well-delimited ostiolar plate divided from metepimeron by deep groovelike metepimeral pseudosuture. Vestibule externally flat, vestibular scar not developed ( Figs 49–50 ). Ostiole ( Figs 47 , 49–51 : o) shifted laterad, situated slightly laterad of meso- and metacetabula (ca. 1/3 of pleuron width), small, round, opening posterolaterad into small periostiolar depression. Peritreme ( Figs 47 , 49–51 : pe, pes) in form of short and narrow spout, slightly depressed; peritremal surface narrow, oriented posteroventrally. Evaporatorium ( Figs 47 , 49–51, 53 : ev) medium-sized, developed on both meso- and metapleuron: on metapleuron covering anteromedian and central portions, reaching ventrally between acetabula, laterally being delimited by sharp longitudinal ridge and posteriorly with groove-like metepimeral pseudosuture, futher projecting laterad as narrow stripe along anterior metapleural margin, reaching nearly end of spiracle; evaporatorium smaller on mesopleuron, occupying posterior margin along spiracle, widened and lobe-like only medially. Surface of evaporatorium punctate, gyrification in form of very fine longitudinal wrinkles. Metathoracic spiracle ( Figs 52–53 : sp) long, narrow, opening ventrally.

Legs ( Fig. 7 ). Femora slightly longer than tibiae, widening from base on, widest anteapically, ventral surface then distinctly narrowed towards apex; oval in cross-section; ventral surface slightly depressed anteapically, unarmed. All tibiae narrow, their superior surface flat, distinctly narrowly emarginated. Tarsi 3-segmented, all segments dorsally rounded.

Pregenital abdomen ( Fig. 7 ). Connexivum exposed ( Figs 6, 8 ), each segment III–VII posterolaterally with low, lobe-like process (widest on segment III, narrowing gradually towards segment VII), connexival outline thus being distinctly undulated. Spiracle of abdominal ventrite II covered by metapleuron. Ventrite III medially and ventrite IV anteromedially with shallow and wide groove to receive labium ( Fig. 7 ). Abdominal venter only slightly convex, medially flattened. Ventrites III–VII laterally with inconspicuous transverse lateral muscle scar (= pseudosuture, i.e., outer impression associated with the inner attachment points of the tergosternal muscle) posteriad of each spiracle; two trichobothria situated transversely posteriad of the muscle scar and posteriad of spiracle (i.e., on spiracular line) on each side of abdomen ( Fig. 55 ).

Male genitalia. Genital capsule ( Figs 56–59 ) trapezoid, wider posteriorly than anteriorly, slightly dorsoventrally flattened, with posterior aperture opening dorsally ( Fig. 57 : pap). Ventral wall deeply depressed posteriorly ( Fig. 58 : vwd), the depression emarginate ventrally by high, wide, rounded, glabrous, transverse ridge ( Fig. 58 : vr), and dorsally by pair of short, narrow, sharp, setose transverse ridges ( Fig. 58 : dr). Ventral rim infolding ( Figs 56–59 : vif) with pair of small depressions basally ( Fig. 58 : vif), then turning anteriad and forming pair of large triangular projections ( Figs 56–57, 59 : vif); dorsal surface of the triangular projections covered with brown setae ( Fig. 57 ), each projection posteromedially with tuft of posteriorly oriented setae ( Fig. 56 ). Lateral rim infolding ( Figs 56–57 : lif) concave, dish-shaped, inner margin posteriorly with narrow incision forming paramere socket ( Fig. 62 ); lateral rim infolding posterolaterally projected into membranous lobe adpressed to the lateral margin of triangular projections of ventral rim infolding ( Figs 56–57, 59, 62 : ml). Dorsal process not developed. Dorsal rim infolding ( Figs 56–57 : dif) short, U-shaped posteriorly, merging with lateral rim infolding. Proctiger cordiform ( Fig. 60 ). Paramere ( Figs 61–63 : p) reduced, weakly sclerotized, long, slender, flattened, C-shaped, devoid of any process; inner margin of paramere in apical half with line of long setae, and with tuft of long setae apically ( Figs 56, 58, 62 ). Phallus ( Figs 64–68 ) robust, phallotheca ( Figs 64–68 : phth) barrell-shaped, provided with a median conical sclerotised process ventrobasally ( Figs 64, 66–68 : vpp). Conjunctiva voluminous, evertible, with a pair of dorsoapical lobes (cp-I?) membranous basally, elongate, hook-like, strongly sclerotized apically; second conjunctival processes (cp-II) sclerotized, hook-like dorsolaterally, ventrolateral portions (median penial plates, Figs 64–68 : mpp) broadened in their distal portions, distal margin provided with 3+3 obtuse denticles, associated with a single ventromedian spoon-shaped process (probably homologous with cp-III). Aedeagus (= vesica sensu authors) strongly curved ventrad at base, then elongate and nearly straight, distal portion weakly curved posteroventrad ( Fig. 68 : aed).

Female genitalia ( Figs 54 , 67–71 ). Laterotergites VIII ( Figs 54 , 67–70 : lt8) subtriangular, dorsally encompassing laterotergites IX and tergite X and medially fused above them. Visible portion of valvifers VIII ( Figs 54 , 67–70 : vf8) subtriangular, dorsally broadly rounded. Laterotergites IX ( Figs 54 , 67–70 : lt9) relatively small, exposed portions kidney-shaped, separated along midline. Valvifers IX ( Figs 54 , 67–70 : vf9) bearing transverse ridge more pronounced laterally. Triangulin well-developed, large, triangular, not visible externally. Gynatrium ( Figs 70–71 ) with pair of fused (pretzel-shaped) ring sclerites ( Figs 70–71 : rs) and a triangular sclerite surrounding spermathecal opening ( Figs 70–71 : so). Spermatheca ( Fig. 71 ): proximal duct ( Fig. 71 : pd) long, distinctly longer than gynatrium, narrow; dilation ( Fig. 71 : dil) long and rather wide in middle, basal constricted portion not developed; distal invagination (= sclerotized rod) ( Fig. 71 : div) subparallel in most of its length, distinctly broadened basally and slightly broadened apically; distal duct ( Fig. 71 : dd) very short, slightly swollen, wider than proximal duct. Intermediate part of spermatheca (= spermathecal pump, Fig. 71 : ip) narrow; proximal flange ( Fig. 71 : pf) narrower than distal flange ( Fig. 71 : df); apical receptacle ( Fig. 71 : ar) hemispherical, low, with three projections directed proximad: the longest projection reaching proximal flange, the intermediate one surpassing distal flange, and the shortest one reaching apex of distal flange.

Differential diagnosis. The new genus can be easily recongized by a combination of its large size; the brick red coloration; the quadrate head without an anteocular spine or incision; the mandibular plates meeting in front of the clypeus; the large and prominent, apically bilobate humeri; and the undulate connexival segments. For detailed comments on its tribal placement see Discussion, for its identification see the Key below.

Etymology. The word Nene is formed as an arbitrary combination of letters, the gender is neuter. CACHAN (1952) named another Triplatygini genus Anoano , which in Malgasy language means a ‘forest insect’ (see KMENT 2012). Coincidentally, Anoano means ‘Yes, Yes!’ in Czech. Because of this I have been tempted since years to accompany Anoano with another generic name, Nene , which means ‘No, No!’ in Czech. In addition, ‘nene’ means a ‘baby’ or, as a term of endearment, ‘darling’ in Portuguese, and in Hawaiian language ‘nēnē’ is the name of the Hawaiian Goose, Branta sandvicensis (Vigors, 1833) (the Czech name of that species being ‘berneška havajská’). This strange opportunity that has been given to me, and it is also a pleasure to dedicate the new name to my charming friend and Czech malacologist Dagmar Říhová (Charles University, Prague), usually known only under her nickname Berneška.

Discussion. The new genus fits well into the tribe Triplatygini as defined by CACHAN (1952), sharing most of the diagnostic characters, especially the shape of the head including large mandibular plates enclosing the clypeus anteriroly, prominent humeri of pronotum, and a spout-shaped peritreme. Based on the recent detailed revisions of all the taxa included in Triplatygini ( KMENT 2008, 2011, 2012; KMENT & BAENA 2015; this paper) the tribe seems to be monophyletic and the following updated diagnosis of the tribe can be given:

Lateral margins of head explanate, sharp. Mandibular plates large, foliaceous, longer than clypeus and meeting in front of it, with anteocular spine or angle and small adjacent anteocular incision (except of Nene ). Antenna pentamerous. Antennal segment I completely covered by head margins or only its apex visible. Humeri of pronotum forming large lobes, apically incised or spinose; lateral margins of pronotum never completely lamellate and sharp. Pronotum and scutellum convex. Scutellum triangular, large, shallowly incised at apices of frena, posteriorly rounded. Costal margin of corium not sharply lamellate. Femora incrassate (especially profemora) but unarmed (except pro- and mesofemora of Tricompastes with few minute tubercles anteapically). Tarsi 3-segmented. Mesosternum grooved, without median longitudinal carina (but low median carina developed in Nene ). Metathoracic scent gland peritreme small, spout-shaped; metapleural evaporatorium small to medium sized. Connexivum large and exposed, lateral margin of abdomen usually undulated or serrate. Base of abdomen without tubercle, spine or groove (except ventrite III grooved in Nene ). Genital capsule with lateral rim infolding well developed, dish-shaped, dorsal processes not developed, and with tendencies of ventral wall to expand dorsally and ventral rim infolding to form triangular, anteriorly oriented projections. Paramere simple, crescent- -shaped ( Tricompastes ) or reduced, devoid of basal process ( Triplatyx ) and desclerotised ( Anoano , Nene ). Phallus with well developed median penial plates (= ventral conjunctival processes). Female genitalia with laterotergites IX small, not meeting each other medially, well developed triangulin and ring sclerites (but ring sclerites rather reduced and plate-like in Anoano and Tricompastes ).

Considering the documented variability of characters, the external morphology and the structure of the female genitalia seem to be of limited information value for the phylogeny of the tribe, being composed of many shared characters, such as symplesiomorphies of Pentatomidae (e.g., the spout-shaped peritreme and possibly also the presence of the triangulin), and putative synapomorphies of Triplatygini as a whole (especially the structure of the mandibular plates and the humeri of the pronotum, though such characters occur, perhaps as parallelisms, also in other tribes of Pentatominae – see e.g. KMENT 2013, or the externally exposed triangulin (in all genera except Nene )), and autapomorphies of the particular genera (e.g., the shape of the humeri in all genera, an orthogonal head and a serrate abdominal margins in Tricompastes , the absence of the anteocular spine, the mesothoracic carina, the labium reaching base of the abdomen, and the ventrite III grooved in Nene , the apical receptacle of spermatheca with many processes in Triplatyx , etc.). To figure out some phylogenetic hypothesis, the structure of the genital capsule and the paramere seem to provide most informative characters. In this regard, Tricompastes seems to be the most different and possibly most distantly related lineage, being characterised by the genital capsule with posterior aperture opening posteriorly, ventral rim infolding small and simple, and large posterolateral projections ( KMENT & BAENA 2015: Figs 21–25 ), and the paramere with a robust and well slerotised body, crescent-shaped, with developed basal process with tuft of setae apically ( KMENT & BAENA 2015: Figs 27–30 ). The three remaining genera share the genital capsule with posterior aperture shifted dorsally, without prominent posterolateral projections but with expanding ventral rim infolding forming a pair of triangular, anteriorly oriented projections ( Figs 56–57 ; KMENT 2008: Figs 35 , 38–39, 41–42 , 44–45 ; KMENT 2012: Figs 17–19 ), and a reduced paramere devoid of basal process ( Figs 61–63 ; KMENT 2008: Figs 47–51 ; KMENT 2012: Figs 17–18 ). Triplatyx is characterised by the posterior aperture oriented posterodorsally, posterolateral angles slightly prominent, the pygophore less gibbose ventrally and the ventral rim infolding less well developed, and the paramere well sclerotised, without setae ( KMENT 2008: Figs 35–46, 47–51 ). Anoano and Nene share the genital capsule somewhat dorsoventrally flattened, with the ventral wall expanding dorsally (and depressed anteapically), the ventral rim infolding with a prominent pair of triangular projections, the posterior aperture opening dorsally, and the paramere reduced and desclerotised but bearing a line of sensory setae.

Figures

Figs 6–8. Nene undulatum sp. nov., habitus of adult (J, paratype from Maroantsetra, body length 14.0 mm): 6 – dorsal view, 7 – ventral view, 8 – lateral view.

Figs 44–47. Nene elongatum sp. nov. 44–45 – head (44 – dorsal view, 45 – lateral view). 46 – humerus of pronotum, dorsal view. 18 – meso- and metapleuron with external scent efferent system of the metathoracic scent gland (evaporatorium dotted).Abbreviations: at – antenniferous tubercle, bc – buccula, ev – evaporatorium, lb1 – labial segment I, lr – labrum, mp – mandibular plate, o – ostiole, pe – peritreme, sp – spiracle. Scale bars: 1 mm.

Figs 48–55. Nene elongatum sp. nov. 48 – head, ventral view (magnification 37×). 49 – meso- and metapleuron with external scent efferent system of the metathoracic scent gland (25×). 50–51 – ostiole and peritreme (50 – 60×, 51 – 230×). 52–53 – metathoracic spiracle (52 – 75×, 53 – 300×). 54 – female external genitalia (30×). 55 – detail of trichobothria and spiracle on ventrite VI (190×). Abbreviations: ev – evaporatorium, o – ostiole, pes – peritremal surface, sp – spiracle, tr – trichobothrium. Scale bars: 0.1 mm – 51, 53, 55; 0.5 mm – 50, 52; 1 mm – 48, 49, 54.

Figs 56–62. Nene elongatum sp. nov. 56–59 – genital capsule: 56 – undissected, dorsal view (magnification 35×); 57 – dissected, dorsal view (35×). 58 – undissected, posterior view (42×); 59 – dissected, lateral view (37×). 60 – proctiger, dorsal view (100×). 61 – paramere, extracted (85×). 62 – paramere, in situ (95×). Abbreviations: dif – dorsal rim infolding, dr – dorsal ridge of ventral wall, lif – lateral rim infolding, ml – membranous lobe of lateral rim infolding, p – paramere, pap – posterior aperture, vif – ventral rim infolding, vr – ventral ridge of ventral wall, vwd – ventral wall depression. Scale bars: 0.2 mm – 60–62; 0.5 mm – 56–59.

Figs 63–68. Nene elongatum sp. nov. 63 – paramere. 64–68 – phallus: 64–66 – uninflated (64 – ventral view, 65 – dorsal view, 66 – lateral view); 67–68 – inflated (67 – ventral view, 68 – lateral view).Abbreviations: aed – aedeagus (= vesica), bp – basal plate, mpp – median penial plates, phth – phallotheca, res – endophallic reservoir, vpp – ventral process of phallotheca. Scale bars: 1 mm.

Figs 69–71. Nene elongatum sp. nov. 69–70 – external female genitalia (69 – intact, 70 – dissected). 71 – spermatheca. Triangulin not depicted. Abbreviations: ar – apical receptacle of spermatheca, dd – distal duct of spermatheca, df – distal flange, dil – spermathecal dilation, div – distal invagination of spermathecal duct, ip – intermediate part of spermatheca, lt8,9 – laterotergite VIII and IX, pd – proximal duct of spermatheca, pf – proximal flange, sp – spiracle, rs – ring sclerite, so – sclerite surrounding spermathecal opening, vf8,9 – valvifer VIII and IX, X – segment X. Scale bars: 1 mm.

Figs 20–26. Jitka elegans sp. nov. 20–24 – genital capsule (20 – undissected, posterior view, magnification 47×; 21 – undissected, postero-dorsal view, 47×; 22 – dissected, posterior view, 45×; 23 – dissected, posteroventral view, 45×; 24 – undissected, lateral view, 47×). 25 – proctiger, dorsal view (120×). 26 – paramere, viewed from above (200×).Abbreviations: app – apical proces of paramere, lif – lateral rim infolding, plp – posterolateral projection of genital capsule, vif – ventral rim infolding. Scale bars: 0.2 mm – 25–26, 0.5 mm – 20–24.

Figs 27–30. Jitka elegans sp. nov.27–30 – paramere (four different views, 150×).Abbreviations:app – apical process of paramere, blp – basal process of paramere, mep – middle process of paramere. Scale bars: 0.2 mm.

Figs 31–35. Jitka elegans sp. nov., phallus. 31–33 – uninflated (31 – dorsal view, 32 – ventral view, 33 – lateral view); 34–35 – partially inflated (34 – ventral view, 35 – lateral view).Abbreviations: aed – aedeagus, bp – basal plate, ds – ductus seminis, mpp – median penial plates, phth – phallotheca, res – endophallic reservoir. Scale bars: 0.5 mm.

Figs 36–38. Jitka elegans sp. nov., female genitalia. 36–37 – external genitalia (36 – intact, 37 – dissected). 38 – gynatrium and spermatheca. Abbreviations: ar – apical receptacle of spermatheca, dd – distal duct of spermatheca, df – distal flange, dil – spermathecal dilation, div – distal invagination of spermathecal duct, ip – intermediate part of spermatheca, lt8,9 – laterotergite VIII and IX, pd – proximal duct of spermatheca, pf – proximal flange, sp – spiracle, rs – ring sclerite, so – sclerite surrounding spermathecal opening, tr – triangulin, vf8,9 – valvifer VIII and IX, X – segment X. Scale bars: 1 mm.

Figs 39–43. Jitka elegans sp. nov., larval dorso-abdominal scent glands (DAG). 39 – larval abdomen in dorsal view (magnification 32×). 40 – anterior pair of DAG (90×). 41 – median pair of DAG (80×). 42 – detail of median DAG (250×). 43 – detail of posterior DAG (200×). Abbreviations: cr – cuticular ridge, es – ear-shaped structure, ev – evaporatorium, o – ostiole covered by valve. Scale bars: 0.1 mm – 42, 43; 0.5 mm – 40, 41; 1 mm – 39.

Figs 14–19. Jitka elegans sp. nov. 14–15 – head of adult (14 – dorsal view, 15 – lateral view). 16–17 – head of larva (16 – dorsal view, 17 – lateral view). 18 – meso- and metapleuron with external scent efferent system of the metathoracic scent gland (evaporatorium dotted). 19 – protibia and protarsus (lateral surface in anterior view of the bug). Abbreviations: at – antenniferous tubercle, bc – buccula, ev – evaporatorium, lb1 – labial segment I, lr – labrum, mp – mandibular plate, o – ostiole, pes – peritremal surface, sp – metathoracic spiracle. Scale bars: 1 mm.