REVISION OF WORLD SPECIES OF PARATELENOMUS DODD (HYMENOPTERA: SCELIONIDAE)

Abstract The genus Paratelenomus Dodd is revised from a worldwide perspective. Three species are described as new: P. angor [Taiwan, Thailand], P. indivisus [Papua New Guinea, Australia], and P. matinalis [Vanuatu]. Paratelenomus bicolor (Dodd) [Australia], P. saccharalis (Dodd) [southern Europe, Africa, tropical Asia, Australia], P. ophiusa (Dodd) [Papua New Guinea, Australia], P. striativentris (Risbec) [Africa, India], and P. tetartus [Indonesia, Malaysia, Philippines] are redescribed. Aphanurus graeffei Kieffer, 1917 and Asolcus minor Watanabe, 1954 are junior synonyms of P. saccharalis (Dodd), 1913. An identification key to species is provided. The relationship of Paratelenomus within Telenominae is discussed; the hypothesized sister group is Nirupama Nixon. Résumé Le genre Paratelenomus Dodd est révisé ici à l’échelle mondiale. Trois espèces nouvelles sont décrites : P. angor [Taiwan, Thaïlande], P. indivisus [Papouasie, Nouvelle-Guinée, Australie] et P. matinalis [Vanuatu]. Paratelenomus bicolor (Dodd) [Australie], P. saccharalis (Dodd) [sud de l’Europe, Afrique, Asie tropicale, Australie], P. ophiusa (Dodd) [Papouasie, Nouvelle-Guinée, Australie], P. striativentris (Risbec) [Afrique, Indes] et P. tetartus [Indonésie, Malaisie, Philippines) sont décrites de nouveau. Aphanurus graeffei Kieffer, 1917 et Asolcus minor Watanabe, 1954 deviennent synonymes récents de P. saccharalis (Dodd), 1913. On trouvera ici une clé d’identification des espèces. La position systématique de Paratelenomus parmi les autres Telenomidae fait l’objet d’une discussion; le groupe soeur est probablement Niruparna Nixon. [Traduit par la Rédaction]


INTRODUCTION
The subfamily Telenominae (Hymenoptera: Platygastroidea, Scelionidae) is a large and unwieldy taxon. Over 800 species have been formally described (Johnson 1992), but this is far from the final total. Attempts have been made to subdivide the two largest genera, Telenomus Haliday and Trissolcus Ashmead, into species groups (e.g. Kozlov and Kononova 1983;Johnson 1984aJohnson , 1984b, but the lineages and their relationships are still far from clear. I have argued that at least one group is easily recognizable and probably monophyletic: the Psix group of genera (Johnson 1988b). This includes Psix Kozlov and Le (see Johnson and Masner 1985), Nirupama Nixon (see Nixon 1935;Johnson 1985), Mudigere Johnson (1988b), and the subject of this revision, Paratelenomus Dodd. Species of Paratelenomus are restricted to the Eastern Hemisphere, from West Africa east to Japan and Vanuatu, and from central Europe and Honshu south to northern Australia and throughout India and Africa. The hosts of these parasitoids are known only for a single species, P. saccharalis (Dodd). It attacks the eggs of bugs of the family Plataspididae (Hemiptera: Heteroptera). Both Paratelenomus and plataspidids are found only in Africa, Asia, Europe, and Australasia. In studying the available material I have been surprised at the relative lack of undescribed species of Paratelenomus. To date, 10 names have been published for the eight species I recognize here. Only three appear to have no available name.
The first species of Paratelenomus was described in 1911, and even though the genus itself was distinguished by Dodd in 1914 (Dodd 1914d), subsequent species have been Description. Head (Figs. 1,3,(5)(6)(7): Hyperoccipital carina absent, occiput rounded onto vertex; frons and lower portion of head with strongly developed fanlike carinae arising near bases of mandibles; central keel arising dorsad of antennal insertions, bifurcating ventrally to pass around antennae, extending dorsad to area of median ocellus, bifurcating there to surround ocellus; orbital carinae arising from near anterior mandibular articulations, extending dorsally along inner orbits to mesa1 edge of lateral ocelli; submedian carinae often present on frons, these arising near anterior mandibular articulation, extending dorsally between central keel and orbital carinae; area below eye between orbital carina and malar sulcus, and surface of gena with fanlike ridges or striae; genal carina well developed, extending dorsad behind eye, sometimes difficult to differentiate from other striae; occipital carina complete, crenulate; clypeus and labrum apparently fused, apex usually medially bidentate; mandibles narrow, sicklelike, without apical dentation, broadly overlapping, apices usually hidden beneath clypeus/labrum; head usually elongate in malar region; eyes sparsely setose; lateral ocelli distinctly separated from inner orbits; preocellar pit absent; Q antennal clava pentamerous, claval formula 1-2-2-2.
Metasoma (Figs. 10,11): First segment, at least, usually more-or-less xanthic, contrasting with dark brown color of mesosoma and remainder of metasoma; T1 without sublateral setae; laterotergite of segment 1 glabrous; T 2 with longitudinal rugulae extending through most of length of tergite, without coriaceous microsculpture between rugulae, smooth and sparsely setose laterally; apex of T2 with narrow smooth band, under high magnification usually revealing extremely fine microsculpture; S2 with extensive arcuate sulci, setal fields rather small, elongate; remaining metasomatic segments with fine microsculpture.

Diagnosis.
Paratelenomus is most closely related to Psix Kozlov and LE and Nirupama Nixon. The unidentate, sicklelike mandibles of Paratelenomus are sufficient to distinguish it immediately, but these are often difficult to see because of the small size of some specimens, particularly when the mandibles are closely appressed within the buccal cavity. In most cases Paratelenomus may quickly be distinguished from Psix by the combination of the xanthic base of the metasoma and the elongate notauli (Figs. 4,12). Species of Nirupama are extremely elongate and the orbital carinae are expressed as sulci; the submedian carinae are absent. The two species Paratelenomus indivisus and Paratelenomus ophiusa both lack notauli and could be confused with Psix. However, they lack the regular rugose-reticulate head and mesonotal sculpture characteristic of Psix, the mandibles are narrow and broadly overlapping, the central keel forks to pass around the median ocellus, and there are no fovea on the posterior side of the meso-metapleural suture.
Discussion. Among telenomines, the sicklelike, unidentate mandibles are found only in Paratelenomus, and support its monophyly. The absence of a line of foveae on the metapleuron flanlung the meso-metapleural suture and the dorsal bifurcation of the central keel are shared with Nirupama and may indicate a sister-group relationship between these two genera. The foveae are present in all other genera of telenomines, often developing into a longitudinal furrow connecting with the metapleural pit. Species of Psix may have the central keel reaching the median ocellus, but at this point it terminates. The two described species of Nirupama also have the base of the metasoma slightly lighter in color than either the following segments or the mesosoma. Nixon (1935) in his original description characterizes the basal segment as reddish-brown. Because we are still unable to postulate confidently a sister group for telenomines the polarization of these character states must be viewed as tentative.
Species of Paratelenomus have the well-developed sulci on the second metasomatic stemite that are also found in the genus Psix (Fig. 11). In Paratelenomus, though, the pattern of the sulci is much more constant, and I have not found them to be useful in discriminating among species. Kozlov and Kononova (1983: 136) place Paratelenomus (as Archiphanurus) in an intermediate position between Trissolcus and Telenomus, and they further claim that it differs from all genera of the subfamily in the sculpture of the frons and gena. The latter assertion is clearly false, as the numerous species of Psix share many of the same facial carinae. I cannot support the former hypothesis as these same features appear to unite Paratelenomus, Psix, and Nirupama. Much has also been made of the shortened postmarginal vein in P. graeffei (= saccharalis). I have found this feature to be very difficult to measure properly and quite variable both intra-and interspecifically. Thus I have not focused on it in this revision.
I had postulated a close relationship among Paratelenomus, Psix, Nirupama, and Mudigere within Telenominae. I am no longer as confident in this hypothesis following this study. All four genera are remarkable for the well-developed carinae on the frons. The first three have the orbital and submedian carinae and the central keel very strongly developed, whereas those of Mudigere are not clearly differentiated. In the meantime I have also become aware of a number of other telenomines, some within Trissolcus and others related to Phanuromyia Dodd, Phlebiaporus Dodd, and Aradoctonus Masner, that also possess these fanlike carinae arising from the mandibles.
L& recently described three Vietnamese species of Paratelenomus (as Archiphanurus): P. aculus, P. irritus, and P. obtusus (L& 1980, 1982). The Institute of Biology in Hanoi is unwilling to mail these specimens and I have been unable to visit so as to examine them. I find the descriptions to contain little of value in recognizing his species. Most of the characters hardly serve to distinguish the taxa from any other species of telenomine, and are of no help in distinguishing among Paratelenomus. One species, P. irritus, is described in the English summary as having no notauli and the first metasomatic tergite brown. This may correspond to the new species described below, P. indivisus, or possibly to P. ophiusa (Dodd). The illustration of P. irritus shows short notauli and a creature more gracile than either of these. (Note that although the caption states that the drawing refers to P. irritus, the figure reference in the text is in the description of P. aculus.) There may be as many as five species of Paratelenomus in Vietnam, and I can find no way to identify P. irritus, P. aculus, and P. obtusus confidently on the basis of the original descriptions.  (Dodd, 1914a) saccharalis (Dodd,19 14 c ) graeffei (Kieffer, 1917) new synonymy minor (Watanabe, 1954) new synonymy striativentris (Risbec, 1950) tetartus (Crawford,191 1) KEY TO WORLD SPECIES OF PARATELENOMUS DODD ( Q )

2'.
Clypeus/labrum clearly much longer than wide, apex rounded medially or drawn to a weak medial point (

Paratelenomus angor new species
Description. Q . Length 1 .lo-1.19 mm (i= 1.13 mm, N = 3). Head: Central keel complete; submedian carinae well developed, running dorsally to mid-height of eye and mingling with sculpture extending ventrally from vertex; lower half of frons between submedian carinae smooth, glabrous; upper half of frons with two to three longitudinal rugae extending from vertex; orbital carina extending from anterior mandibular articulation along inner orbit to mesal edge of lateral ocellus; area between orbital and submedian carinae in upper half of frons with single longitudinal ruga; area below eye between orbital carina and malar sulcus with four to five ridges radiating from anterior mandibular articulation; a single carina present between malar sulcus and genal carina; clypeus/labrum pentagonal, apex bidentate. Mesosoma: Dark, concolorous with head; notauli well developed, nearly percurrent; disk of mesoscutum strongly coriaceous throughout, nearly pustulate; transscutal articulation noticeably narrowed medially, crenulae along posterior margin also narrower medially than laterally, but not so markedly, these define a transverse fusiform surface on scutellum, this with strong raised coriaceous microsculpture similar to mesoscutum; mesopleural carina present, flanked anteriorly by three to five deep foveae; space between fore and mid coxae bridged by costae from coxal margins; anteroventral portion of mesepisternum otherwise largely smooth, glabrous; acetabular field very small, circular, finely coriaceous; episternal foveae indistinguishable, foveae along mesopleural carina merging with those flanking pronotal-mesepisternal suture; mesopleural scrobe longitudinally costate, grooves not reaching mesopleural carina; mesepimeron indicated as a convex fusiform surface anterior to meso-metapleural suture; metapleural triangle well developed; metapleuron without crenulae flanking meso-metapleural suture. Metasoma: First segment dark brown, concolorous with mesosoma and remaining segments of metasoma; T1 with three lateral setae; apex of T2 with band of extremely fine microsculpture.  submedian carinae abbreviated, represented by low ridge extending from anterior mandibular articulation to only below mid-height of eye, weakly defined; orbital carina complete, extending from mandibular articulation to lateral ocellus; frons medially smooth, glabrous, in upper third with four to five low, weakly meandering longitudinal mgae; area between orbital carina and malar sulcus below eye with three to five fanlike ridges radiating upward from anterior mandibular articulation toward eye; gena with two ridges between malar sulcus and genal carina; crenulae arising from occipital carina short; gena near dorsal end of genal carina finely coriaceous; clypeus/labmm pentagonal, apex bidentate. Mesosoma: Golden yellow, contrasting with dark head; notauli present, nearly percurrent; mesoscutum with finely pebbled surface microsculpture; transscutal articulation narrowed medially; crenulae on posterior margin of scutellum narrowed medially, disk of scutellum coriaceous; mesopleural carina absent, without foveae marking its course; area between fore and mid coxae bridged by crenulae flanking coxal cavities; anteroventral portion of mesepistemum largely smooth; acetabular field small, finely coriaceous; epistemal foveae not differentiated; mesopleural scrobe longitudinally sulcate; mesepimeron not differentiated; metapleural triangle well developed; metapleuron without crenulae flanking meso-metapleural suture. Metasoma: Extensively xanthic, central areas of T2, S2, and apex of metasoma dark brown, dark pigmentation gradually developing, without marked transition; T1 with two lateral setae; apex of T2 smooth. 0 . Unknown.

Distribution. Northern Australia.
Diagnosis. This species is immediately recognizable by its striking coloration. It is otherwise very similar to P. saccharalis in that the submedian carinae are poorly developed and the mesopleural carina is not indicated. In xanthic species such as this the details of microsculpture are very difficult to see under a light microscope because the cuticle is almost translucent.  , v-vi.1982, 10.vi.1982, 16.vii.1982, 24.vii. 1982,2.xi. 1982,30.xi. 1982,79  Etymology. The name indivisus refers to the lack of division of the disk of the mesoscutum by notauli.

Distribution. Northem Australia and Papua New Guinea.
Diagnosis. Only two species of Paratelenomus lack notauli in some form, P. indivisus and P. ophiusa. This species is distinguished from P. ophiusa by the entirely dark metasoma, and the bidentate apex of the clypeusPabrum.

Paratelenomus matinalis new species
Description. Q . Length 0.80-0.99 mm (T= 0.90 mm, N = 38). Head (Fig. 3): Central keel complete, extending dorsad from bifurcation above antennal insertions to median ocellus; submedian carinae well developed, extending dorsad to merge with vertexial sculpture; lower half of frons between submedian carinae glabrous, smooth or with strongly effaced coriaceous microsculpture; upper half of frons with three to four rugulae parallel to central keel; orbital carina extending to mesa1 margins of lateral ocellus; area between orbital and submedian carinae above level of upper half of eye with setal bases strongly pustulate; area below eye between orbital carina and malar sulcus with three deep grooves defining four ridges; a single groove present posterior to malar sulcus and simultaneously flanking genal carina, thus a single ridge between malar sulcus and genal carina; crenulae arising from occipital carina extending to genal sulcus; clypeus/labrum pentagonal, apex bidentate. Mesosoma (Fig. 4): Notauli present, sinuate, nearly percurrent; mesoscutum between notauli weakly rugose-reticulate, with coriaceous microsculpture, setal bases strongly pustulate; sculpture effaced laterad of notauli, practically absent entirely near scutellum; transscutal sulcus strongly narrowed medially, crenulae within barely visible along midline of body; crenulae bordering posterior margin much shorter medially than laterally; crenulae and transscutal articulation define transverse fusiform surface on scutellum, this with fine coriaceous microsculpture laterally, nearly smooth medially, setal bases weakly pustulate; mesopleural carina present, flanked by two to four foveae at level of acetabular field of microsculpture; space between fore and mid coxae bridged by elongate crenulae flanking coxal cavities; anteroventral portion of mesepistemum occupied by expanded crenulae from coxal cavities and acetabular carina; acetabular field very small, oval, finely coriaceous; episternal foveae indistinguishable, anterior edge of mesopleural scrobe abutting crenulae flanking pronotal-mesepistemal suture, scrobe entirely transversely sulcate; two postepimeral foveae below fore wing base; metepisternal triangle well defined; mesometapleural suture without crenulae posteriorly. Metasoma: First segment bright goldenyellow, contrasting with following dark brown segments; T1 with two lateral setae; apex of T2 with narrow band of extremely fine microsculpture. Etymology. The name matinalis refers to the eastem distribution of this species in relation to the others in the genus.

Distribution. Islands of Vanuatu (formerly New Hebrides).
Diagnosis. This species, beautiful for its crisply defined sculpture and color pattern, shares with P. angor and P. striativentris the presence of a mesopleural carina. From those two it may be distinguished by the lack of mesoscutal microsculpture laterad of the notauli. The contrast in color between the first metasomatic segment and the rest of the body is particularly striking.

Distribution.
Probably widespread in southern Palearctic, Africa, tropical Asia, and northern Australia.
Diagnosis. This is by far the most common and most widespread species of Paratelenomus. It is generally characterized by its small size and delicate sculpture. Paratelenomus bicolor is very similar, but quite different in coloration. Paratelenomus tetartus also has welldeveloped notauli and lacks the mesopleural canna. Paratelenomus saccharalis may be distinguished from P. tetartus by the pentagonal, bidentate clypeusPabrum. The short submedian carinae on the frons is fairly characteristic of P. saccharalis through much of its range, but in specimens from Japan and Korea the carinae continue dorsally to merge with the vertexial sculpture.
Remarks. Through most of its range Paratelenomus saccharalis is bisexual, but in Europe the species is unisexual (Bin and Colazza 1988). Wall (1928) published a brief account of some aspects of the biology of this species, referring to it as a chalcid. Later (Wall 1931) he referred to it as P. tetartus (determined by A.B. Gahan). Some of the material from that study is now deposited in OSUC.
The male paralectotype is no longer protected beneath the cover slip with the female. I chose not to try to remount it because it is already broken and because of its age.

Diagnosis.
Other species in this genus with notauli present and the foveae marking the edge of the mesopleural carina are l? matinalis and P. angor. Paratelenomus striativentris is distinguished from them by the completely sculptured mesoscutum and scutellum, the multiple carinae in the position of the submedian carinae, and the light base of the metasoma.