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Matriz de caracteres As marcas ‘?’ na matriz indicam quando um estado não pôde ser

determinado. A mesma marca quando em frente ao nome de uma espécie indica incerteza em sua identificação. Táxons precedidos por ‘EX’ são grupos externos.

CAPÍTULO 2: Estudo das relações filogenéticas supraespecíficas de Hybothoracini Bouček (Hymenoptera; Chalcididae)

Revision of Steninvreia Bouček (Hymenoptera; Chalcididae) BRUNO CANCIAN DE ARAUJO1,2

1 Programa de Pós-Graduação em Ciências Biológicas, Universidade Federal do Espírito Santo, Av. Fernando Ferrari, nº 514,

, Vitória, Espírito Santo, Brazil, 29.975-910 . 2 chalcididae@gmail.com

Abstract

The genus Steninvreia is revised, diagnosed and compared with other related Hybothoracini. One new synonym is proposed: Steninvreia noyesi Narendran n. syn. under Steninvreia anupama Narendran The four valid species (S. anupama Narendran,

S. edgari Bouček, S. petiolata Bouček and S. tricarinata (Girault)) are described and

illustrated. Five types are analyzed (four holotypes and one paratype). A key for all species is provided.

Key words: Haltichellinae, Chalcidoidea, taxonomy, Australasia.

Introduction

Steninvreia was proposed by Bouček (1988) based on the type species Xenarretocera tricarinata (Girault, 1927) and on two other new species, S. edgari and S. petiolata, all

from Australia. Narendran (1989b) combined the type species of Thanushyama (T.

anupama Narendran, 1989a, from Philippines) with Steninvreia and described S. noyesi

from India, totalizing five species into the later genus.

The species are small, 2.6-3.9 mm long, with a great range of morphological forms (Bouček, 1988). All species have a bright black color, except for the antenna and apex of the leg segments testaceous. Despites the comparison with Proconura made by Narendran (1989a), the genus is more related with Notaspidium, as cited by Narendran (1989b), which shares, among other features, short body pilosity, smaller than the diameter of the associated fovea, and the presence of the one lobe in the base of the anterior wing. According to Bouček (1988) the sexual dimorphism is small, once some males present shorter and more stout antenna, more convex scutellum and last gastral segment truncated. Host data are unknown.

While revising Hybothoracini, tribe which the genus belong, we accessed the type material of all described species of Steninvreia, leading us to realize that the genus cam have a better morphological delimitation and the species need a revision, with better descriptions and illustrations. Thus, this paper attempts to compile data about

Steninvreia, propose a better morphological definition for the genus, revise the number

of valid species and improve the quality of descriptions and illustrations of the species, as well as proposing the first identification key to all valid species.

Material

The follow acronyms are used for collections containing type material on which this study is based (with the names of individuals who assisted in the loan of material given in parenthesis) or other collections cited.

ANIC - Australian National Insect Collection, Camberra, Australia; DZCU - Calcutta University, Calcutta, India;

QMBA - Queensland Museum, Queensland, Australia. (Susan Wright). SAMA - South Australian Museum, Adelaide, Australia. (Peter Hudson).

USNM - United States National Museum of Natural History, Washington, DC, USA (Michael Gates).

Methods

Morphological analysis and measurements were made with a stereo microscope Leica M205C fitted with a 5 mm/100 divisions ocular grid, with an illumination system Leica LED 5000 RL. A piece of translucent mylar acetate was used between the led and specimen to reduce glare. The measurements were taken in different magnification and expressed in millimeters. The terminology of structures and sculptures follow Bouček (1988), except by propodeum sublateral carina, the longitudinal carina between the submedian carinae and plicae, and propodeum lateral panel, the lateral area posterior to the metapleuron. The terms foveate and punctate follow Harris (1979). The follow acronyms refer to the morphological terms of measurements: DAO, diameter of the anterior ocellus; DW, dorsal width of the eye; ES, length of the stigmal vein; F, funicular segment (exemple: F1 first, F2 second); FV, width of the frontovertex; FW, frontal width of the eye; H, height; L, length; M, length of the marginal vein; ME (malar

space), distance between the ocular and oral margins; OOL, (ocell-ocular line) small distance between lateral ocellus and the adjacent eye; OVP, length of the ovipositor sheath; POL, (post-ocellar line) distance between hind ocelli; S, sternite of the gaster (exemple: S1 first, S2 second); SM, length of the submarginal vein; T, gastral tergites (exemple:, T1, first: T2 second); TCS (torulus-clypeus space), distance between the inferior margin of the antennal socket and the clypeus margin; W, width; WE, width of the eye in frontal view; WPE, width of the parascrobal space at the level of the scrobal width; WS, largest width of the scrobe.

The morfological characteristics were filled into a matrix and translated to descriptions with software DELTA (Dallwitz et al., 1993). As the examined material is restricted to types, all this section was written verbatim in the caput of the species, with additions between brackets. Image capture were made by an automated system Leica Z16 APO and LAS software, illuminated by a multichannel L.E.D. dome system with 120 super white LED’s developed by the first author. This system uses indirect diffuse light to avoid glare. The multifocus images was combined and retouched in Helicon Focus 6.

Results

Steninvreia Bouček, 1988

Steninvreia Bouček, 1988: 58-60. Type species Xenarretocera tricarinata Girault, 1927 Tanushyama Narendran 1989a: 46. Type species Tanushyama anupama Narendran,

1989a

Diagnosis. Body glossy black except for apex of the segments of the antenna and legs

testaceous or yellow; Body usually cylindrical with short pilosity, shorter than the associated fovea. Head globose to triangular in frontal view, usually with a angle above the scrobe leaving the vertex in a more vertical plan; occiput deep, minimum as deep as wide in dorsal view; scrobe narrow, shallow and poorly delimited, weakly reticulate to polished; antenna short, scape reaching maximum a half of the eye high; flagellum short, smaller than double of the scape length, with a conical or wedged clava; interantennal projection very short or absent; malar carina absent; genal carina very strong. Pronotum subsquare with parallel sides in dorsal view, narrower than mesoscutum at the tegula level, exposing a wide prepectus; scutellum more or less flat and subcircular, never projected over the dorsellum and metanotum, which are always

visible in dorsal view; metafemur with a narrow base and a shot comb of teeth, restrict to the distal ventral third, ending in a small dentiform projection; metatibia short, medially curved with a clavate apex; propodeum usually long and subcilindric. Gaster with a variable shape, usually narrow anteriorlly; petiole very short, visible only ventrally, or long, but always with an irregularly foveate integument; T1 engraved- reticulate with conspicuous median and lateral carinae, which never reaches a half of the tergite.

Distribution. Australia (Australia Capital Territory, Queensland, New South Wales,

South Australia), India (Kerala), Philippines (Mindanao). Bouček (1988) mentioned four undescribed species for Australia, one for Africa, one for the south Asia, three for New Guinea and one for Salomons, which shows a predominant Australasian distribution.

Biology. Unknown.

Discussion. Steninvreia belongs to the group of Hybothoracini species with S1 sides not

smooth, more related with the genus Tropimeris and Notaspidium which shares short body pilosity, smaller than the diameter of the associated fovea. Despites shares with

Tropimeris a extreme reduction or absence of the interantennal projection, it’s closer to Notaspidium which shares, among other features, the presence of the one lobe in the

base of the anterior wing. Differs of this by a prepectus anteriorly broad (due to a subsquare pronotum with parallel sides in dorsal view, narrower than the mesoscutum at the level of the tegula), absence of the metallic bright and absence of the foiciform membranous area in the internal face of the metacoxa. Except for S. edgari which has a dorsoventral flattening body, the other species have a lanky and subcylindrical body, a feature that names the genus, with globose head and deep occiput. The mesopleurom venter is proportionally broader than others Hybothoracini (except one species of

Notaspidium), always longer than wide and covered by a short and dense pilosity. The

scutellum is usually flat and subcircular, never projected over the dorsellum and metanotum, which are always visible. The propodeum is long and subcilindric, except in S. edgari wherein is short with the lateroposterior angles projected posteriorly involving the anterior region of T1, as in Hybothorax. The metafemur has a narrow base with a short comb of teeth restrict to the distal third and starting in an inconspicuous toothform projection. The metatibia is always short, medially curved with the apex wider than the base, with a club aspect. The petiole could be very short, visible only in dorsal view, or long as in S. petiolata, but always with an integument covered by a

irregular furrows. The gaster has a variable shape, longer in S. petiolata and S.

tricarinata, shorter in S. edgari, usually ending truncate. Bouček (1988) and Narendran

(1989b) described a greenish bright in some species of the genus, but we didn’t realize this feature (a tendency to green) under diffuse or direct white light, even using the methodology described in Aguiar (2005). The cylindrical bare body and the information known about the hosts of the related genus (e.g. Notaspidium) may suggest a concealed host as a leafminer or a stick/seed borer.

Key to the species of the Steninvreia Bouček of the world

1. Petiole more than 1.5x as long as high (fig 12); propodeum as long as wide; gaster very long, 2.83x as long as wide (fig 11) ... S. petiolata Bouček, 1988 - Petiole maximum as long as high (figs 4, 13); propodeum wider than long;

gaster maximum 2.36x as long as wide (fig 15) ... 2 2. Head more than 1.59x as wide as high, with a conspicuous subtriangular aspect (fig 6); propodeum with lateroposterior region headmost than its center (fig 8); S1 anteriorly foveate (fig 5) ... S. edgari Bouček, 1988 - Head less than 1.25x as wide as high, usually with a globose aspect; propodeum with lateroposterior region rearmost than its center (figs 3, 11 ); S1 anteriorly carinate ... 3 3. Mesepisternum with foveate convex area behind the prepectus, T1 almost as

high as T2 in lateral view (fig 13) ... S. tricarinata (Girault, 1927) - Mesepisternum with smooth triangular flat area behind the prepectus (fig 2), T1 much higher than T2 in lateral view (fig 4) ... S. anupama (Narendran, 1989)

Steninvreia anupama (Narendran, 1989b)

(Figs 1-4)

Tanushyama anupama Narendran, 1989a: 46; Holotype male (USNM, examined). Steninvreia anupama (Narendran, 1989b): 165.

Steninvreia noyesi Narendran, 1989b: 165; Holotype female [QMBA (not DZCU as

Type material. T. anupama: HOLOTYPE (male, USNM) “[PHILIPPINES] Mindanao

Zamboanga; L F Baker Collection 1927.”; point-mounted, head, mid leg, right front wing, posterior tarsi and gaster glued on card; missing parts: antenna after scape, pronotum and fore legs. S. noyesi: HOLOTYPE (female, QMBA) “INDIA: Kerala. C. U. Campus; NARENDRAN Party April 1987; pl 9637; QM Reg. No. T57888”; point- mounted, in good condition.

Description. Female. lenght = 3.64 mm. Body glossy black except for scape, pedicel,

fore and mid leg except coxae, base of the metafemur, apex of the metatibia and metatarsus testaceus. Head almost as wide as high. Vertex, in frontal view, with slightly elevated margins. Anterior ocellus distant 2.30x its own diameter from the scrobe (fig 2); scrobe rugulose; APL regular, 1.13x DOA. Frons projected making it in different plans the vertex and inferior face; torulus-ocular carina absent; antennal socket separated by 0.70x your own diameter, in lateral view, not reaching the lateral limit of the oral fossae; scape reaching maximum the middle of the eyes, narrow, 6.50x as long as wide, ventral face convex, pilose; funicular pilosity less than 1/5 the segment width, sensilla exceeding the distal limit of the funicle; clava symmetric. Clipeus angulate; labrum concave; mandible with a basal depression; malar space very short, 3.13x DOA; genal carina wide. Measurements: Head L 0.48, H 0.72, W 0.77; FV 0.22; LS 0.28; WPE 0.10; TCS 0.03; ME 0.25; DAO 0.08; APL 0.09; OOL 0.02; POL 0.15; Eye H 0.41, DW 0.25, FW 0.22; Scape L 0.39, W 0.06; Pedicel L 0.09, W 0.06; F1 L 0.03, W 0.05; F2 L 0.04, W 0.07; F3 L 0.05, W 0.08; F4 L 0.05, W 0.08; F5 L 0.06, W 0.09; F6 L 0.06, W 0.10; F7 L 0.06, W 0.10; F8 L 0.06, W 0.11; Clava L 0.20, W 0.12.

Pronotum and mesoscutum foveae separated by your own diameter or more. Pronotum rounded (fig 3), lateral panel flat, without fovea (figs 1, 2). Tegula with sparse pilosity; mesepisternum delimited by strong pleural suture, with smooth triangular flat area behind the prepectus (fig 2); epcnemial carina medially interrupted, mesopleural venter with posterior margin not elevated. Wing marginal vein glabrous, median fold line conspicuos, posterior margin arched, glabrous. Scutellum flat, interstice engraved-reticulate; frenal carina conspicuous. Metapleuron interstices polished, ventral projection present. Metafemur comb of teeth lobulate; metatibia slightly arched, ventral, median and external internal carinae absent; metapulvillus with flat apex, minumum as wide as the base; base of the tarsal claw forming a lobe. Propodeum short, 0.59x as long as wide, sublateral carinae strong and rectilinear,

lateroposterior region pilose, rearmost than the center of propodeum (figs 3, 4), lateroventral projection present, lateral panel subrectangular. Measurements. Pronotum L 0.45, W 0.64; Mesoscutum L 0.27, W 0.70; Scutellum L 0.41, W 0.50; Propodeum L 0.44, W 0.75; Posterior wing L 1.61, W 0.66, SM 0.64, M 0.09; Metacoxa L 0.39; Metafemur L 0.56, W 0.29.

Gaster. Twice as long as wide; petiole maximum as long as high, venter projected anteriorly; T1 much less high than T2, tightly convex (fig 4), engraved reticulated, median and submedian carinae conspicuous, lateral and adjunct carinae absent, lateral polished and glabrous; S1 carinate, ventral tooth and ventral carinae absent; T2 reticulate, dorsal posterior margin tightly curved (fig 3); T6, in lateral view, convex and pilose; espiracle with low borders; T7+8 without a laminar carinae covering to the base of the cerci. Measurements. Gaster L 1.11, W 0.55; T1 L 0.64; T7+8 L 0.05, W 0.07; OVP 0.05.

Male. Lenght ~ 4.55 mm. Similar to female except as follows. Anterior ocellus

distant 2.00x its own diameter from the scrobe, scrobe reticulate. Antennal socket separating by your own diameter. Clipeus fused. Tegula with pilosity concentrated on the border. Fore wing marginal vein with 3 long bristles in the dorsal face. Scutellum more convex than in female. T1 dorsally with a more weak reticulation, T2 dorsal posterior margin slightly curved.

Distribution. India (Kerala), Philippines (Mindanao). Biology. Unknown.

Discussion. We propose here Steninvreia noyesi as a junior synonymous of S. anupama

oncethe differences between these specimens are sexual dimorphism. Otherwise, both of them were composed by one specimen of each sex, with a close geographical distribution. This species is morphologically close to the S. petiolata, sharing globose head as wide as high, scrobe in a more inclined plane than the vertex (less accentuated than in S. petiolata), a deeper occiput in dorsal view and a rounded pronotum. It is easy to recognize by the mesepisternum delimited by strong pleural suture, forming smooth triangular flat area behind the prepectus and by a tightly convex T1 much less high than T2.

Steninvreia edgari Bouček, 1988

(Figs 5-8)

Steninvreia edgari Bouček, 1988: 59; Holotype male (ANIC), Paratype male (SAMA,

examined), other 2 Paratypes male (SAMA).

Type material. PARATYPE (male, SAMA) “Between Bower & Florieton; 14. Aug.

1959, S[outh] A[ustralia]; C. F. Gross & P. Aitiken”. Condition: Point-mounted, entire except for anterior right leg and tarsus lost.

Description. Male. Length = 4.24 mm. Color glossy black except for antenna, tegula,

fore and mid leg except coxae, apex of the metacoxa, border of the metafemur, metatibia and metatarsus testaceus. Head wide, 1.59x as wide as high (fig 6). Vertex, in frontal view, with elevated margins. Anterior ocellus distant 1.20x its own diameter from the scrobe, scrobe reticulate; APL very long, 3.20x DOA. Frons not projected; torulus-ocular carina absent; antennal socket separated by 2.90x your own diameter, in lateral view, not reaching the lateral limit of the oral fossae; scape long, surpassing the middle of the eyes (fig 6), stout, 4.75x as long as wide, ventral face flat to slightly convex, glabrous; pilosity of the funicles more than 1/3 of the segment width, funicle sensilla exceeding the distal limit of the funicle; clava wedged. Clipeus fused; labrum flat; mandible without a basal depression; malar space very long, 7.20x DOA; genal carina narrow. Measurements: Head L 0.60, H 0.85, W 1.35; FV 0.63; LS 0.30; WPE 0.11; TCS 0.00; ME 0.36; DAO 0.05; APL 0.16; OOL 0.04; POL 0.31; Eye H 0.59, DW 0.39, FW 0.38; Scape L 0.57, W 0.12; Pedicel L 0.22, W 0.09; F1 L 0.07, W 0.10; F2 L 0.09, W 0.11; F3 L 0.08, W 0.11; F4 L 0.09, W 0.12; F5 L 0.09, W 0.12; F6 L 0.09, W 0.13; F7 L 0.07, W 0.14; F8 L 0.08, W 0.15; Clava L 0.24, W 0.15.

Pronotum and mesoscutum foveae separated by less than your own diameter (fig 7). Pronotum subsquare, lateral panel convex, foveate (fig 5). Tegula with pilosity concentrated on the border; mesepisternum delimited by a strong pleural suture, with a foveate area behind the prepectus; epcnemial carina continuous. Wing marginal vein with 2 conspicuous bristles in the dorsal face, median fold line conspicuos, posterior margin retilineous, pilose. Scutellum convex, interstices engraved-reticulate; frenal carina inconspicuous (fig 8). Metapleuron interstices engraved-reticulate, ventral projection present. Metafemur comb of teeth almost rectilineous; metatibia tightly arched, ventral internal, median and external carinae incomplete. Propodeum very short,

0.35x as long as wide, sublateral carinae weak and sinuous, lateroposterior region glabrous, headmost than the center of propodeum (fig 8), lateroventral projection absent, lateral panel subtriangular. Measurements. Pronotum L 0.57, W 1.12; Mesoscutum L 0.63, W 1.25; Scutellum L 0.63, W 0.99; Propodeum L 0.44, W 1.25; Posterior wing L 2.25, W 0.90, SM 0.78, M 0.14; Metacoxa L 0.60; Metafemur L 0.96, W 0.57.

Gaster short, 1.27x as long as wide; petiole higher than long, venter not projected anteriorly; T1 slightly less high than T2, almost flat, punctate, median and submedian carinae inconspicuous, lateral and adjunct carinae present, lateral punctate, pilose; S1 foveate (fig 5), ventral tooth and ventral carinae present; T2 foveate, dorsal posterior margin slightly curved; T6, in lateral view, convex and pilose; espiracle with low borders; T7+8 with a laminar carinae covering the base of the cerci. Measurements. Gaster L 1.27, W 1.00; T1 L 0.75; T7+8 L 0.03, W 0.14.

Fenale. Unknown.

Distribution. Australia (Australia Capital Territory, South Australia). Biology. Unknown.

Discussion. This is the most peculiar species of the genus. It shares triangular head and

subsquare pronotum with S. tricarinata, but the head have a more triangular aspect, 1.59x as wide as high. The body have a flat dorsoventral aspect, with a gaster 1.27x as long as wide versus a minimum twice as long as wide of the other species. The propodeum is very short, 0.35x as long as wide, only in the genus with the lateroposterior region headmost than the center of propodeum and with a T2 tightly foveate.

Steninvreia petiolata Bouček, 1988

(Figs 9-12)

Steninvreia petiolata Bouček, 1988: 59; Holotype female (USNM, examined), Paratype

male (BMNH).

Type material. HOLOTYPE (female, USNM) “Illawara; N[ew] S[outh] Wales; H.

Petersen”. Condition: Point-mounted, entire except for posterior right tarsus lost.

Description. Female. Length = 4.40 mm. Color Glossy black except for apex of the

elevated margins. Anterior ocellus distant 3.20x its own diameter from the scrobe, scrobe reticulate; APL regular, 1.13x DOA. Frons tightly projected making it in different plans the vertex and inferior face (fig 10); torulus-ocular carina absent; Antennal socket separated by 1.10x your own diameter, in lateral view, not reaching the lateral limit of the oral fossae; scape short, not reaching the middle of the eyes (fig 10), stout, 4.86x as long as wide, ventral face flat to slightly convex, glabrous; pilosity of the funicles more than 1/3 of the segment width, funicle sensilla exceeding the distal limit of the funicle; clava symmetric. Clipeus fused; labrum flat; malar space short, 3.38x DOA; genal carina narrow. Measurements: Head L 0.48, H 0.76, W 0.76; FV 0.31; LS 0.21; WPE 0.11; TCS 0.01; ME 0.27; DAO 0.08; APL 0.09; OOL 0.01; POL 0.15; Eye H 0.47, DW 0.23, FW 0.23; Scape L 0.34, W 0.07; Pedicel L 0.10, W 0.06; F1 L 0.04, W 0.06; F2 L 0.06, W 0.08; F3 L 0.07, W 0.09; F4 L 0.07, W 0.09; F5 L 0.07, W 0.09; F6 L 0.07, W 0.10; F7 L 0.07, W 0.10; F8 L 0.08, W 0.11; Clava L 0.20, W 0.11.

Pronotum and mesoscutum foveae separated by less than your own diameter. Pronotum rounded (fig 11), lateral panel flat, foveate. Tegula with sparse pilosity; mesepisternum delimited by a strong pleural suture, with a foveate reticulated area behind the prepectus; epcnemial carina continuous, venter of the mesopleuron with a posterior margin elevated. Wing marginal vein with 3 conspicuous bristles in the dorsal face, median fold line conspicuos, posterior margin arched (fig 9), pilose. Scutellum convex, interstices polished; frenal carina very strong (fig 11). Metapleuron interstices polished, central projection present. Metafemur comb of teeth almost rectilineous; metatibia tightly arched, ventral internal and median carina absent, external carina incomplete; metapulvillus with flat apex, minumum as wide as the base; base of the tarsal claw forming an angle with a pair of setae. Propodeum as long as wide, sublateral carinae strong and rectilinear, lateroposterior region pilose, rearmost than the center of propodeum, lateroventral projection absent, lateral panel subrectangular. Measurements. Pronotum L 0.45, W 0.68; Mesoscutum L 0.51, W 0.77; Scutellum L 0.47, W 0.55; Propodeum L 0.57, W 0.58; Posterior wing L 2.12, W 0.75, SM 0.85, M 0.10; Metacoxa L 0.25; Metafemur L 0.61, W 0.27.

Gaster very long, 2.83x as long as wide (fig 11); petiole 1.5x as long as high, venter projected anteriorly (fig 12); T1 almost as high as T2, convex, engraved reticulated, median carina inconspicuous, submedian carinae conspicuous, lateral carinae present, adjunct carinae absent, lateral slightly reticulate and pilose; S1 carinate, ventral tooth and ventral carinae absent; T2 reticulate, dorsal posterior margin slightly

curved (fig 11); T6, in lateral view, convex and pilose; espiracle with low borders; T7+8 without a laminar carinae covering the base of the cerci. Measurements. Gaster L 1.78, W 0.63; T1 L 0.76; T7+8 L 0.02, W 0.05; OVP 0.04.

Male. According Bouček (1988) It’s very similar to female in color, body and

most of its details, even in antennae, only scapus slightly stouter. Also propodeum,

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