Archaeoteleia

Early, J. W., Masner, L. & Johnson, N. F., 2007, Revision of Archaeoteleia Masner (Hymenoptera: Platygastroidea, Scelionidae)., Zootaxa 1655, pp. 1-48 : 3-8

ARCHAEOTELEIA MASNER

Archaeoteleia Masner 1968: 652. Type: Archaeoteleia novaezealandiae Masner , by original designation. Original description, key to species, systematic position. Masner 1976: 5, 12 (description, keyed); Johnson 1992: 344 (catalog of world species); Austin & Field 1997: 10, 68 (structure of ovipositor system, discussion of phylogenetic relationships, genus misplaced in Sparasionini ).

Description: Medium to large, length 2.2-6.4 mm, usually slender; legs, wings, antennae often rather elongate, flightless females more robust, with shorter appendages; usually rather pale-colored, yellow to brown, sometimes brown to very dark brown (some with pale regions actually light green when alive), female antenna sometimes with A4 or A4+A5 gradually or distinctly lighter in color than adjacent antennomeres; body sculpture varying from smooth to densely punctate or rugulose, shining to matte; usually macropterous, but with females in some species brachypterous, micropterous or apterous; external sexual dimorphism apparent in structure of antenna, T1, apex of metasoma.

Head slightly transverse when viewed dorsally; vertex rounded, hyperoccipital carina absent; occipital carina present, complete medially; lateral ocellus distinctly separated from inner orbit by distance of at least 1-1.5 ocellar diameters, OOL greater than or equal to LOL; eye moderate to large in size, glabrous or with short setae; frons convex, without frontal depression, median longitudinal carina present; head with large raised interantennal process between toruli; torulus opening laterally on interantennal process; submedian carina absent; orbital carina present, arising at anterior mandibular articulation, continuing dorsally to lateral ocellus; lower frons with dense fanlike striae; interocular space variable in width, from subequal to distinctly less than height of eye; inner orbits very weakly divergent ventrally, subparallel; clypeus roughly triangular to subpentagonal in outline, high, usually with prominent angulate lateral corners, convex, postclypeus and anteclypeus not differentiated, apical margin usually straight; malar sulcus present, continuous with posterior orbit of eye, often obscured among fanlike facial striae; gena moderately widened, convex; labrum not exposed; mandible strong, deeply bidentate, teeth transversely oriented, subequal in size; maxillary palpus 5-segmented, penultimate segment cylindrical; labial palpus 3-segmented; antenna 12-merous in both sexes; radicle differentiated from base of A1, inserted apically into A1 at a distinct angle to longitudinal axis of A1; basal portion of A1 distinctly curved medially; apex of female antenna with gradually widened, cylindrical clava composed of conical A12, subquadrate A6-A11, apically expanded A5; gustatory sensilla on female antenna arranged in longitudinal pairs on apical antennomeres; claval formula A5-A12/1-2-2-2-2-2-2-1 or A6-A12/ (1,2)-2-2-2-2-2-1; A3 always longest flagellomere, usually longer than A1, always longer than length of A2; male antenna with tyloid position variable, either on A4, A5, A4+A5, A4+A5+A6, tyloid arising at base of segments; male flagellomeres usually extremely elongate, with short pubescence.

Mesosoma in dorsal view longer than wide, in lateral view usually high, usually convex dorsally; pronotum in dorsal view with pronotal humeral carina and well-developed lateral shoulders, epomial corners distinct, often sharply defined forming strong angle or upward projection; vertical epomial carina usually absent or very weak; lateral face of pronotum weakly concave, forming shallow scrobe for reception of fore leg; netrion usually clearly delimited anteriorly by arc of deep foveae, very broad, as wide as tegula, open ventrally; anterior margin of mesoscutum meeting pronotum anteriorly, mesoscutum with arched anterior margin, narrowed behind tegulae; admedian lines sometimes well-developed; parapsidal line almost always present; notaulus variably developed, complete, abbreviated, or entirely absent; skaphion absent; transscutal articulation flanked posteriorly by well developed line of long crenulae; scutellum transverse, often constricted or impressed along midline, posterolateral corners usually protruding in form of sharp spine or hook, sometimes unarmed; axilla large, subtriangular, posterior margin foveolate, without raised flange; metanotum fairly broad, unarmed, dorsellum bounded by distinct line of foveae dorsally and ventrally; dorsal surface of propodeum with moderate pilosity, usually excavate medially in female to house horn on T1, excavate in male of one species, sometimes with weak submedian longitudinal keels, these sometimes expanded into small, weak tooth; mesopleuron large, prominent; mesopleural depression well-developed; mesopleural carina moderately developed to absent, lower end of carina extending to posterior margin of mesopleuron; sternaulus absent; mesopleural pit present; anterior margin of ventral portion of mesepisternum straight, not protruding between fore coxae; mesepisternum and mesepimeron separated by line of well-developed foveae; episternal foveae absent; dorsal corner of mesepimeron not produced into tooth or hook; anteroventral portion of metapleuron rounded, not separated from lateral face by carina; metapleural pit absent; posterior margin of metapleuron not produced; posterolateral corners of propodeum not projecting posteriorly; legs usually long, slender; hind leg, particularly coxa, elongate, length of hind coxa twice length of midcoxa, posterior surface not striate; femora usually not incrassate; trochantellus present on all legs; outer surface of tibia without spines; tibial spur formula 1-2-2, outer spur shorter than inner on mid, hind leg; hind tibia without longitudinal keels; tarsal formula 5-5-5; tarsomeres tapering in width apically; hind tarsus usually cylindrical, not laterally compressed; pretarsal claws simple; apex of fore wing usually reaching or surpassing apex of metasoma, largely clear, in some species with cloud of pigmentation basally between R1 (marginal vein) and r-rs (stigmal vein), marginal cilia short; R (submarginal vein) straight, broadly separated from costal margin, extending through basal half of length of fore wing, subapically with distinct bulla, forked apically beyond bulla, usually with moderate setae, rarely with large stiff dark bristles; R1 extending to costal margin forming marginal vein, in some species continuing apically along costal margin to form postmarginal vein; postmarginal vein variable from short to several times longer than stigmal vein; membrane of wing deeply pigmented between bulla and point where R1 reaches costal margin, forming pseudostigma, pseudostigma relatively short and wide in Chilean species, long and narrow in New Zealand species; r-rs longer than R1, slightly downturned apically, angle between R1, r-rs nearly perpendicular in species from New Zealand, usually acute in species from Chile; no other tracheate veins in fore wing; apical portion of fore wing with pigmented lines in position of Rs, M, Cu; hind wing with R tracheate only in its basal half, never reaching hamuli and costal margin; R without bristles; 3-4 hamuli present; cilia on posterior margin of hind wing subequal in length with marginal cilia of fore wing.

Metasoma usually elongate, weakly depressed, sutures between segments usually constricted to form undulating profile; submarginal ridge well-developed, laterotergites and laterosternites narrow, very weakly sclerotized, hyaline; female with 6 terga and sterna visible externally, segments 1-4 subequal in length; male with 8 terga and 7 sterna visible externally, segments 1-5 subequal in length, segments 6-8 gradually shortening distally, apical segments minute; female T1 with distinct horn; S1 with weak median longitudinal keel, not laterally compressed, not extending anteriorly between hind coxae; anterior margin of S2 straight; sterna without felt fields; female with base of segment 6 much narrower than apex of segment 5; female T6 with apical margin convex, without median raised field of microsetae or secretion; apical tergite of male with distinct, short, cylindrical cerci, slightly longer than wide; female with T7+T8 broadly separated from T6 by membranous telescopic tube (i.e., Scelio-type ovipositor), tube with two segments, basal segment strongly pigmented.

Diagnosis: Archaeoteleia is distinguished from most other scelionids by the 1-2-2 tibial spur formula. It shares this plesiomorphic character state with the genera Sparasion Latreille, Sceliomorpha Ashmead, Nixonia Masner, and Neuroscelio Dodd. The distinguishing characters are summarized in the following key.

1 Malar sulcus present; head with strong fanlike facial striae reaching orbits of eyes; laterotergites and laterosternites hyaline; female with ovipositor horn on T1........................................................ Archaeoteleia

- Malar sulcus absent; head without fanlike facial striae; laterotergites and laterosternites well sclerotized; female T1 without ovipositor horn .............................................................................................................. 2

2 Antenna 14-segmented; paired gustatory sensilla of female antennal clava arranged side by side. Nixonia

- Antenna 12-segmented; paired gustatory sensilla of female antennal clava arranged longitudinally..........3

3 Pronotum without sharp dorsal transverse carina; body short, squat........................................ Neuroscelio

- Pronotum with sharp transverse carina extending from epomial carina on one side of the body to the other; body elongate, cylindrical............................................................................................................................. 4

4 Radicle not differentiated from remainder of scape; base of scape U-shaped, inserting on the underside of the interantennal prominence.................................................................................................. Sceliomorpha

- Radicle clearly differentiated by a transverse line and distinctly narrower diameter, inserted into base of scape at a distinct angle to the longitudinal axis of the scape; antenna arising frontolaterally from interantennal prominence........................................................................................................................ Sparasion

Geographic Distribution: New Zealand and the Valdivian part of Chile.

Link to Distribution Map. [http://atbi.biosci.ohio-state.edu:210/hymenoptera/eol_scelionidae.content _page?page_level=3&page_id=taxon_page_data&page_version=453&page_option1=M]

Biology: Two New Zealand species parasitize the eggs of Gymnoplectron spp. (Orthoptera: Rhaphidophoridae , Macropathinae ).

Relationships: Ever since its original description (Masner 1968), Archaeoteleia has been considered to be a plesiomorphic group within the Scelionidae . Species possess a number of plesiomorphic character states, including paired tibial spurs on the mid and hind tibiae; maxillary palpus with 5 segments and labial palpus with 3 segments; the radial vein in the fore wing has a distinct break, or bulla, near its apex; and the antennal radicle is attached to the remainder of the scape at a sharp angle. In the first modern attempt to group scelionine genera into tribes, Kozlov (1970) failed to mention Archaeoteleia . Largely on the basis of the plesiomorphic characters, Masner (1976) placed Archaeoteleia in the tribe Sparasionini together with Sparasion , Sceliomorpha , and the fossil genus Electroteleia Brues (from Baltic amber). Kozlov and Kononova (1990) considered the monotypic tribe Nixoniini to be the most plesiomorphic extant group in the family, but in their cladogram they postulated a sister-group relationship between Nixoniini and Sparasionini . This hypothesis was based upon the proposition that in both tribes T6 and T7 are fused in the female metasoma, whereas in the male T7 and T8 are fused. In fact, this character state is not shared among all Sparasionini .

Austin & Field (1997) extensively surveyed the structure of the ovipositor system and apical metasomatic segments in Scelionidae and Platygastridae . They reported, surprisingly, that the putatively plesiomorphic genus Archaeoteleia possesses the complex apomorphic Scelio-type ovipositor. In this system, the ovipositor is extruded from the apex of the metasoma by internal hydrostatic pressure, the sclerotized portions of the ovipositor (gonocoxae, gonapophyses, and gonoplacs) are borne at the apex of a membranous telescopic tube, and the composite sclerite T7+T8 is broadly separated from T6 and extruded from the end of the metasoma when the ovipositor is in use. Austin & Field considered the shared possession of the complex of characters that constitute the Scelio-type ovipositor to represent strong evidence for a novel monophyletic group within Scelionidae which they called the Scelionini sensu lato. They then suggested, because it retains a number of plesiomorphic characters, that Archaeoteleia should probably be placed in a tribe separate from other Scelionini .

From this new perspective, a number of characters also support the removal of Archaeoteleia from the rest of the Sparasionini . Archaeoteleia has a strong set of fanlike carinae that arise from the anterior mandibular articulation; a fine, but well-developed malar sulcus; the labrum is small and hidden beneath the clypeus; and, in all species but one, the lateral corners of the clypeus are strongly expanded. These characters are shared with many of the so-called more "apomorphic" genera of scelionids, but are lacking in Sceliomorpha , Sparasion , Nixonia , Neuroscelio , and almost all members of the family Platygastridae .

In dried specimens of Archaeoteleia the terga are wider than the sterna and the specimens appear to have the classical scelionid submarginal ridge. Typically this is composed of narrow laterotergites that flex beneath the central tergum and articulate with an impressed submarginal groove on the corresponding sternum. In most Scelioninae , all Teleasinae , and many Platygastridae there is also a narrow, differentiated, and articulated laterosternite on each segment. In the past it has been supposed that this complex lateral articulation of the metasomatic segments may be associated with the production of hydrostatic pressure for extrusion of the ovipositor (e.g., Austin etal. 2005). However, although we believe that they are present, the laterotergites and laterosternites in Archaeoteleia are extremely lightly sclerotized and nearly unpigmented. Their presence is only indicated by a different surface texture, most clearly seen in specimens that have the metasoma bloated and have been critical-point dried, and in scanning electron micrographs, by a fold along the lateral margin of each segment. This suggests that extrusion of the tubular Scelio-type ovipositor is not dependent upon the presence of a laterotergite/laterosternite locking system in the metasoma.

Murphy et al. (2007) recently examined the relationships among genera within Platygastroidea on the basis of three genes (18S, 28S, COI). They reported that Archaeoteleia groups together with Neuroscelio Dodd and, in some analyses, with Sparasion , together forming the sister group of all other Platygastroidea. These results suggested, again, that the Scelionidae is paraphyletic (as in Austin & Field 1997), but also that the tubular ovipositor observed in Archaeoteleia was independently derived from the remainder of the Scelionini sensu lato. If these results are corroborated, then this will further strengthen the proposal that Archaeoteleia should be classified separately from other Scelionini .

Key to species of Archaeoteleia

The New Zealand and South American species form two distinct groups. Those from New Zealand have the scutellar spines small, weak, or absent (e.g., Fig. 23); the pseudostigma is elongate and narrow, not triangular (e.g., Figs. 1, 8, 20, 25); and the stigmal vein forms an angle of greater than 45° with the postmarginal vein. All known species are macropterous. In contrast, the Chilean species have well-developed scutellar spines that are very prominent in brachypterous forms (Figs. 36, 58, 72); the pseudostigma is short, broad and triangular (Figs. 31, 39, 61); and the stigmal vein forms an angle of less than 45° with the postmarginal vein. The species of these two groups are keyed out separately.

NEW ZEALAND SPECIES (females of A. waipoua n.sp. unknown)

1 Posterior corners of scutellum with short pointed tubercle or spine (Figs. 4, 11, 19, 27; fore wing with distinct pigmented area under marginal vein between bulla and stigmal vein (Figs. 8, 16, 25) ............... 2

- Posterior corners of scutellum rounded, without any trace of tubercle or spine (Figs. 15, 23); pigmented area under marginal vein between bulla and stigmal vein very faint or absent ....................................... 5

2 Scutellum barely bilobate (Fig. 27); metasomatic tergites coarsely rugose (Fig. 29); mid and hind basitarsi strongly compressed laterally (Fig. 24); clypeus narrow, rounded, without sharply angulate anterolateral corners (Fig. 28); female unknown ..................................................................... A. waipoua n.sp.

- Scutellum bilobate (Figs. 4, 11, 19); metasomatic tergites with fine longitudinal coriarious sculpture (Figs. 2); all basitarsi normal, cylindrical (Figs. 1, 8, 16); clypeus with sharply angulate anterolateral corners (Figs. 5, 9, 17) .............................................................................................................................. 3

3 Female T1 horn smooth, shining (Fig. 19); transverse pronotal carina straight, with prominent, angulate epomial corners (Fig. 19); notauli ending anteriorly behind epomial corners; pronotal shoulders narrow, maximum width about 0.5 times width of tegula; scutellum usually concolorous with mesoscutum, never clearly paler.......................................................................................... A. novaezealandiae Masner

- Female T1 horn sculptured, finely coriarious (Figs. 4, 11); transverse pronotal carina curved around anterior margin of mesoscutum, epomial corners distinct but not prominent; notauli (or, when incomplete, their projected trajectory) ending at or between epomial corners (Figs. 4, 11); pronotal shoulders broad, at least 0.75 times width of tegula; scutellum usually distinctly paler than mesoscutum (Figs. 8, 11).............................................................................................................................................................4

4 Female: body length 3.4-4.0 mm; metasoma 3.4-3.7 times longer than wide; T6 2.3-2.6 times longer than wide. Male: body length 3.1-3.7 mm; antenna with A5 carinate........................... A. gilbertae n.sp.

- Female: body length 5.0-5.6 mm; metasoma 4.2-4.6 times longer than wide; T6 3.0-3.5 times longer than wide. Male: body length 4.4-4.7 mm; antenna with A5 or A4+A5 or A4-A6 carinate (Figs. 6, 7)... .................................................................................................................................... A. chambersi n.sp.

5 Notauli complete (Fig. 23); mesopleuron below mesopleural depression covered with dense, whitish, appressed pilosity (Fig. 22); female T1 horn sculptured; body length 4.8-6.4 mm ....... A. onamata n.sp.

- Notauli abbreviated anteriorly (Fig. 15); mesopleuron below mesopleural depression almost bare (Fig. 14); female T1 horn smooth, shining; body length 2.5-3.7 mm ....................................... A. karere n.sp.

SOUTH AMERICAN SPECIES

Females

1 Micropterous or apterous: fore wings at most a minute scale not extending beyond metanotum (Figs. 36, 58, 72)...............................................................................................................................................2

- Macropterous: fore wings extending at least to apex of metasoma (Figs. 30, 42, 46, 50, 54, 64, 68) .................... 4

2 T6 about as long as wide, length 0.8-1.1 times maximum width (Fig. 59); T1 horn in lateral view short, almost vertical, not reaching level of mesoscutum, remote from scutellar spines (Fig. 58)...................... ..................................................................................................................................... A. pygmea Masner

- T6 at least twice as long as wide (Figs. 37, 72); T1 horn slanting forward, reaching above level of mesoscutum, projecting between base of scutellar spines (Figs. 36, 74)................................................3

3 T2-T5 with paired longitudinal depressions (Fig. 37); A1 slightly longer than A3; notauli complete; epomial corners not projecting upward............................................................................. A. dispar n.sp.

- T2-T5 without paired longitudinal depression; A1 at least 2 times as long as A3; notauli absent; epomial corners projecting upwards, appearing like small spines in lateral view (Fig. 74)............................ ................................................................................................................................. A. submetallica n.sp.

4 Length of A1 about 2 times length of A3; mesoscutum almost flat in lateral view (Fig. 66); T1 horn entirely sculptured, with median longitudinal carina on posterior face (Fig. 67); submarginal vein with very prominent, upright, strong black bristles; femora distinctly incrassate .................. A. robusta n.sp.

- A1 shorter than A3; mesoscutum convex in lateral view; T1 horn smooth, not carinate; submarginal vein with fine pale setae; femora not incrassate ...................................................................................... 5

5 Mesoscutum entirely evenly sculptured, punctate or finely coriaceous, without smooth areas (Figs. 49, 53, 57) ..................................................................................................................................................... 6

- Mesoscutum with some smooth areas (Figs. 33, 45, 71) ........................................................................8

6 T6 short, about as long as wide; posterior margin of T5 straight; T1 horn small, weakly developed (Figs. 50, 53); mesopleuron entirely sculptured (Fig. 52) ............................................................ A. penai n.sp.

- T6 elongate, about 3 times as long as wide; posterior margin of T5 concave (Figs. 46, 54); T1 horn well developed, reaching level of mesoscutum (Figs. 48, 54); upper part of mesopleuron smooth (Figs. 48, 56) ........................................................................................................................................................... 7

7 Upper frons, vertex, occiput entirely punctate, sculpture similar to that of mesoscutum (Figs. 55, 57); A6, A7 paler than remaining segments of clava (Fig. 54) ......................................... A. puncticeps n.sp.

- Upper frons with longitudinal sculpture, becoming weaker towards ocelli (Fig. 47); most of vertex and occiput usually with transverse wrinkles, differing from sculpture of mesoscutum (Fig. 49); clava uniformly dark (Fig. 46).................................................................................................... A. mellea Masner

8 T1 with low hump, completely longitudinally costate (Figs. 68, 70, 71); length of A3 2.7 times length of A4 .................................................................................................................................. A. simulans n.sp.

- T1 with horn completely smooth in upper part (Figs. 35, 44, 45); length of A3 1.5 times length of A4 ....................... 9

9 Body entirely black to very dark brown (Figs. 30-35); posterior two-thirds of median lobe of mesoscutum, scutellum predominantly glabrous (Fig. 33).................................................... A. araucana Masner

- Body brownish or yellowish (Figs. 42-45); entire median lobe of mesoscutum, scutellum with setigerous punctures (Fig. 45)................................................................................................. A. gracilis Masner

Males (males of A. penai n.sp., A. submetallica n.sp. unknown)

1 Postmarginal vein with well-developed tubular section that is longer than stigmal vein.............................2

- Postmarginal vein at most with rudimentary, short tubular section that is shorter than stigmal vein, often obscured by thick, pigmented pseudostigma ............................................................................................... 4

2 Notauli present (Fig. 39); A4, A5 carinate basally; netrion well defined, indicated by row of foveolae .....................3

- Notauli absent (Fig. 61); only A4 with small basal carina; netrion not well defined................. A. pygmea Masner

3 A1 longer than A3; A6-A9 straight, cylindrical; anterior margin of T1 sharply pointed, fitting into wedge-shaped excavation of propodeum......................................................................................... A. robusta n.sp.

- A1 shorter than A3; A6-A9 arcuate; anterior margin of T1 normal, propodeum without excavation.......... ............................................................................................................................................... A. dispar n.sp.

4 Mesoscutum entirely evenly sculptured, punctate or coriaceous, without smooth areas............................5

- Mesoscutum with some smooth, polished areas..........................................................................................6

5 A4, A5 carinate; upper frons, vertex, occiput entirely punctate, similar to sculpture of mesoscutum........... ....................................................................................................................................... A. puncticeps n.sp.

- A4 carinate, A5 rarely with rudimentary carina; upper frons with longitudinal sculpture, this becoming weaker towards ocelli; most of vertex, occiput smooth and shining, differing from sculpture of mesoscutum .................................................................................................................................... A. mellea Masner

6 Only A4 with keel; body predominantly light-colored, yellowish brown........................ A. gracilis Masner

- Both A4 and A5 with keels; body distinctly darker, dark brown to black....................................................7

7 Posterior two-thirds of mesoscutum glabrous.............................................................. A. araucana Masner

- Entire mesoscutum with setigerous punctures................................................................... A. simulans n.sp.