THE NEARCTIC SPECIES OF HOLOTELEIA KIEFFER (HYMENOPTERA: SCELIONIDAE)

Abstract The Nearctic species of the genus Holoteleia are revised. Two named species are redescribed and five new species are described: H. armigera from Florida; H. coriacea, H. laticeps, and H. polita from eastern Canada and the United States; and H. elegans, widespread east of the Rocky Mountains. A diagnosis of Holoteleia and a key to the Nearctic species are given. Potential impact of environmental degradation on the frequency of Holoteleia species in North America is discussed. Résumé Les espèces néarctiques du genre Holoteleia ont fait l’objet d’une révision. Deux espèces connues sont décrites de nouveau et cinq nouvelles espèces sont décrites : H. armigera de Floride, H. coriacea, H. laticeps et H. polita de l’est du Canada et des États-Unis, et H. elegans, très répandue à l’est des Rocheuses. On trouvera ici la diagnose du genre Holoteleia, ainsi qu’une clé d’identification des espèces néarctiques. L’influence que peut avoir la détérioration de l’environnement sur la fréquence des espèces d’Holoteleia en Amérique du Nord est examinée. [Traduit par la Rédaction]


INTRODUCTION
The seven species of the genus Holoteleia Kieffer in America north of Mexico comprise minute (1.4-1.8 mm), spindle-shaped, elongate, yellowish-orange wasps, often with shortened wings. Only two species were known prior to this revision (Muesebeck 1979); five species are described as new to science. This increase in the number of species reflects the general situation in faunal exploration of most groups of parasitic Hymenoptera in North America. More effort from entomologists and greater support by granting agencies are urgently needed to stimulate the faunal surveys. Environmental degradation and global climatic change are new challenges to all concerned biologists.
The geographic distribution of Holoteleia in its present concept (Masner 1 9 8 0~) is restricted to the Holarctic super-region. Holoteleia nigriceps (Kieffer) is the only known species in the western Palearctic, recorded from Italy and France (Kieffer 1908(Kieffer , 1926, Czechoslovakia (Masner 1956), and Austria (Masner 1976). The seven Nearctic species presently recognized are predominantly distributed in the eastern part of North America (Figs. 1-7), with only H. bicolor (Hanington) extending considerably westward to Alberta (Fig. 1). This pattern corresponds in general to the geographic distribution of ground crickets (Gryllidae, Nemobiinae) in North America (Daniel Otte, personal communication).
The biology and behaviour of species of Holoteleia are little known. The wasps are presumed to parasitize eggs of various ground crickets (Gryllidae, Nemobiinae). Even though no Nearctic species has been reared from cricket eggs, all seven species were frequently caught in traps together with nymphal or adult-ground crickets especially during the late summer and fall. Severin (1935) described the biology of a closely related scelionid wasp, Calliscelio marlattii (Ashmead), an egg parasitoid of Gryllus assimilis Fabricius, a serious pest of agricultural crops in South Dakota. Up to 50% of all eggs of the pest were reportedly parasitized by the above wasp. Therefore, the two frequent and widely distributed Nearctic species of Holoteleia, viz. bicolor and parvipennis (Melander and Brues), could play a significant role in controlling ground crickets in habitats such as meadows. Furthermore, the two above-mentioned species seem to prefer generally manmade or man-altered situations such as fields, gardens, lawns, etc. In contrast, the remaining five Nearctic species of Holoteleia are markedly less frequent, preferring natural habitats such as peat bogs, sand dunes, or forests. Pan trapping is the most productive collecting method. Six species have a distinct late summerearly fall peak of occurrence; only H. armigera is clearly vernal in Florida.
Faunal surveys conducted by the author during the past 20 years in the Ottawa-Hull region showed a remarkable decrease in populations of two local common species, viz.
H. bicolor and H. pawipennis. Pan traps operated in 10 locations in the Ottawa-Hull area, representing four major ecosystems (urban meadows, marsh, sands, forest), yielded gradually fewer individuals during [1970][1971][1972][1973][1974][1975][1976][1977][1978][1979][1980][1981][1982][1983][1984][1985][1986][1987][1988]. A near total crash in populations occurred in [1989][1990], with only a few individuals caught during the entire season. A similar decline was monitored on Nearctic species of Duta Nix. (Masner 1991) and other genera of local scelionid wasps. Altogether 48 species representing nine genera were monitored; thousands of specimens were examined and tabulated. It is interesting to note that the only European member of Holoteleia (H. nigriceps Kieff.) is very rare in European collections. During the past years I was able to examine only a dozen individuals of the above species (Masner 1976). The more advanced environmental degradation in Europe may be a possible explanation for the rarity of H. nigriceps Kieffer. Nearly all of the 1029 specimens of Nearctic Holoteleia examined in this study are deposited in the Canadian National Collection of Insects and Arachnids (CNCI) and were collected in the past 2 decades. The deposition of specimens other than in CNCI is stated.
Metasoma elongate, spindle-like, narrowed at base; laterotergites narrow, submarginal ridge well impressed; T1 and T2 with longitudinal costae, T1 in female usually with hump, hump not developed in H. coriacea and H. polita; T3 largest segment of metasoma, smooth or sculptured; T6 in female subtriangular, not attenuate; T7 in female attached to and extruded with ovipositor.
In North America the concept of Holoteleia was precisely defined recently (Masner 1980a) to include only two eastern Nearctic species: H. bicolor (Harrington) and H. parvipennis (Melander and Brues). These were previously classified in Baryconus auct. (Hamngton 1899;Muesebeck and Walkley 195 I), Caloteleia Ashmead (Melander and Brues 1903), Ceratoteleia Kieffer (Muesebeck and Walkley 195 I), Calotelea Westwood (Muesebeck and Masner 1967). Members of Holoteleia can be distinguished from those of Calotelea by the non-striate cheek, by the notaulus being at least partly developed (in most species), by the non-modified dorsellum, and by the absence of a skaphion on the mesoscutum. From Calliscelio Ashmead (= Ceratoteleia) the members of Holoteleia differ by the non-modified dorsellum and by the subtriangular metasomal tergite 6 in females. In contrast to all Nearctic species of Calotelea (Masner 1980b) and Calliscelio, the Holoteleia species in North America frequently display facultative brachyptery or are only brachypterous.
Holoteleia belongs to the tribe Calliscelionini of the subfamily Scelioninae (Masner 1976). The female ovipositor is housed internally in a membranous tube which is telescoped out during oviposition (Figs. 21,22); abdominal tergites 8 and 9 (apparent metasomatic tergite 7) are extruded at the tip of the telescoped tube, similarly as in females of the tribe Scelionini (Field and Austin 1994 -T3 almost entirely smooth; T2 with costae not exceeding basal half of tergite ( Q ) (Fig. 12 Holoteleia armigera sp.nov. (Figs. 1,19,20) Diagnosis and Recognition ( 9 0 ) . Frons entirely densely coriaceous with scattered deep setigerous punctures, setae extremely short, appressed; notaulus not developed; scutellum entirely coriaceous; wings reduced to narrow strips without venation, not exceeding TI; mesopleural carina ventrally with parallel chain of foveolae; upper apices of propodeal keels sharply pointed, projecting; female T1 anteriorly with hump; T2 with extremely short keels in basal fifth of tergite; T3 entirely finely coriaceous, with dense appressed yellow pilosity. Holoteleia armigera is unique among Nearctic species in having pointed and sharply projecting upper apices of the propodeal keels, extremely short keels on T2, the notaulus absent, and narrow strip-like wings without venation in both sexes.
Mesosoma in dorsal view longer than wide (58:43); mesoscutum evenly densely coriaceous, with scattered appressed yellowish hairs; notaulus not developed; humeral and suprahumeral sulci well impressed, non-foveolate; tegula reduced but present; transscutal suture without foveolae at side, narrow and deep across its entire course; scutellum entirely finely coriaceous, with short appressed yellowish hairs, without crenulae along posterior margin (in front of scutellar rim); dorsellum with ventral lip very narrow; propodeum anterodorsally deeply excavate with anterior apices sharply pointed and projecting bladelike, posterolateral comer of propodeum sharply pointed, sides of propodeum (below spiracle) with fine dense pilosity; mesosoma in lateral view convex, longer than high (58:44); side of pronotum entirely finely coriaceous; netrion relatively narrow with deep foveolae along anterior margin, foveolae larger than half netrion width; mesopleural depression smooth and glabrous; mesopleural carina complete, sharp, flanked ventrally by row of irregular foveolae and generally fine rugulosity (Fig. 20); mesepisternum below metapleural carina entirely finely coriaceous with scattered pilosity; acetabular carina sharp, nonfoveolate; mesepimeron divided from mesepistemum by row of foveolae, foveolae becoming indistinct in lower part; metapleuron smooth and glabrous except for patch of sculpture above hind coxa, with row of fine foveolae along anterior margin and with deep non-foveolate median sulcus; fore wing narrow, strip-like without veins and bristles, reaching to anterior margin of TI; hind wing similar to fore wing, slightly exceeding posterior margin of propodeum.

Diagnosis and Recognition
Holoteleia bicolor can be distinguished from the sympatric H. parvipennis by the presence of foveolae or irregular rugae below the mesopleural carina, also by the longer longitudinal costae on T2 and the distinct coriaceous sculpture on T3. From H. coriacea, it differs primarily by the relatively smoother sculpture on the frons, presence of a hump on TI, and the generally larger size.
Biology. Host and habits unknown. Adults occur in open, sunny habitats such as grassland and bogs, with peak occurrence in late summer and early fall (August-September).
Variation. The fore wing in females (frequently) and males (rarely) may be shortened, rather narrow, reaching only to about the middle of T3. Foveolae under the mesopleural carina may appear in some individuals as irregular impressions or (rarely) may be almost obliterated. For this reason H. bicolor appears twice in the key (cf. p. 78). The coriaceous sculpture on T3 may be effaced in the middle part of the tergite, usually better expressed at sides. The hump on T1 in females may vary slightly in degree of development. The anterior half of the midlobe of the mesoscutum may rarely bear a dark spot similar to individuals of H. elegans.
Remarks. Harrington's (1899) type series consists of mixed individuals of H. bicolor and H. pawipennis (Sarazin 1986). Holoteleia bicolor is very close to the Palearctic H. nigriceps Kieffer from which it differs principally by the sculpture of the lower frons and that of T3; in H. bicolor the lower frons is almost entirely smooth and T3 is finely coriaceous with smoother middle part, whereas in H. nigriceps the lower frons has strong transverse wrinkles and T3 is distinctly longitudinally striate in its middle part.

Diagnosis and Recognition
Holoteleia coriacea is unique among Nearctic species principally because of the entirely coriaceous frons. The absence of a hump in T1 in the female and the short strip-like wings in both sexes further characterize the species. Members are also the smallest among the Nearctic species of Holoteleia.
Mesosoma in dorsal view longer than wide (47:35); mesoscutum evenly densely scaly-like coriaceous with dense appressed hairs; notaulus complete, deeply incised particularly in posterior half; humeral and suprahumeral sulci deep, non-foveolate; scutellum anteriorly with 2-3 deep foveolae and deep transscutal suture medially; scutellar disc with dense coriaceous sculpture and few setigerous punctures especially in anterior half, sculpture becoming delicate in posterior margin, posterior margin of scutellum (in front of scutellar rim) with fine foveolae; dorsellum with narrow smooth ventral lip; propodeal keels meeting below dorsellum, anterodorsal apices of keels not pointed and not projecting; posterolateral comer of propodeum sharply produced into point; mesosoma in lateral view convex dorsally, longer than high (47:37); side of pronotum smooth medially with patch of dense microsculpture in front of spiracle; netrion narrow, smooth, foveolate along anterior margin, foveolae smaller than netrion width; mesopleural depression smooth and glabrous; mesopleural carina strong, complete, with parallel row of deep foveolae ventrally; mesepistemurn below mesopleural carina mostly smooth, with scattered hairs; acetabular carina sharp, not foveolate ventrally; mesepimeron divided from mesepistemum by row of deep foveolae; metapleuron smooth and glabrous with row of foveolae along anterior margin and with deep non-foveolate median sulcus; fore wing shortened, narrow strip-like, only slightly wider than width of hind femur, not exceeding posterior margin of T3, without black bristles on submarginal vein; marginal vein longer than short stigma1 vein; postmarginal vein slightly longer than marginal vein; basal and medial veins not indicated.

Biology.
Host and habits unknown. The adult wasps seem to prefer open, wet habitats, such as bogs, fens, and marshes.
Variation. The fore wing in the female may very rarely attain full size, widely surpassing the tip of the metasoma; fore wing in the male may vary from reaching the posterior margin of T3 to slightly surpassing the tip of the metasoma.
Etymology. Coriacea derived from coriaceus in Latin meaning leather-like, referring to the sculpture of the head, especially the frons.
Holoteleia ekgans sp.nov. (Figs. 4,12) Diagnosis and Recognition ( Q a ) . Upper part of frons with even fine coriaceous sculpture and scattered pilosity, only lower part of frons smooth; scutellum entirely coriaceous; mesopleural carina ventrally with parallel chain of foveolae; upper apices of propodeal keel not pointed and not projecting, female T1 with smooth hump; longitudinal keels on T2 not exceeding basal half of tergite; T3 almost entirely smooth; mesoscutum usually bicoloured, orange with large dark spot on anterior half of midlobe and smaller darker spots on lateral lobes.
Holoteleia elegans can be distinguished from H. bicolor by the smooth T3, by the costae on T2 not exceeding the basal half of the tergite, by having the upper part of A1 and entire A2 dark brown, and by the distinct large circular dark spot on the anterior half of the mesoscutum.
Description. FEMALE (Holotype). Length 1.7 mm. Colour predominantly orange-yellow, with head almost black, anterior half of A1 and rest of antenna dark brown to black, mesoscutum with large dark circular spot in anterior half of midlobe and with smaller darker spots on lateral lobes, scutellum, hump on TI, most of T2, posterior part of T3, and T4-T7 dark brown, tegula and metapleuron light brown, labrum, palpi, mandible, radicle, basal half of Al, pleura, posterior half of mesoscutum, anterior half of T3, and legs including coxae orange-yellow; wings slightly infuscate.
Mesosoma in dorsal view longer than wide (50:40), cervical part of pronotum not foveolate; mesoscutum evenly densely reticulate with scattered setigerous punctures, with dense decumbent yellowish pilosity, notaulus incomplete, deeply incised in posterior two-thirds of mesoscutum; humeral and suprahumeral sulci deep, non-foveolate; scutellum entirely and evenly coriaceous, with few scattered setigerous punctures; posterior margin of scutellum (in front of scutellar rim) distinctly foveolate; dorsellum with ventral lip well developed; propodeum (below dorsellum) partly excavate with anterodorsal points noncontiguous, posterior margin of propodeum carinate, not projecting to sharp points; mesosoma in lateral view longer than high (50:41); side of pronotum smooth medially, with patch of microgranular sculpture in front of spiracle and with fine rugulose sculpture along dorsal margin; netrion smooth, foveolate along anterior margin, foveolae about half netrion width; mesopleural depression smooth and glabrous; mesopleural carina complete, sharp, flanked ventrally by row of deep foveolae; mesepistemum below mesopleural carina partly smooth, with fine scattered hairs and with fine sculpture near acetabular carina; acetabular carina not foveolate; mesepimeron divided from mesepisternum by row of deep foveolae; metapleuron smooth and glabrous with row of deep foveolae along anterior margin and with deep nonfoveolate sulcus; fore wing attenuate, not exceeding tip of metasoma with 9 semierect bristles along submarginal vein, marginal vein shorter than stigmal vein (5:7), postmarginal vein longer than marginal vein (175); basal and median veins non-pigmented.

Material
Remarks. The males of H. elegans have the longitudinal costae on T2 exceeding the basal half of the tergite. In this respect they resemble males of H. bicolor but differ from them by having T3 almost entirely smooth.
Etymology. Elegans derived from the latin elegans meaning fine or beautiful, referring to the fine combination of two colours on the mesoscutum. Holoteleia laticeps sp.nov. (Figs. 5,14,18) Diagnosis and Recognition ( Q 0 ) . Head in dorsal view strongly transverse; upper half of frons (including area in front of anterior ocellus) densely coriaceous, with scattered setigerous punctures; scutellum entirely sculptured; mesopleural carina ventrally without foveolae; upper apices of propodeal keels closely approximated, not pointed, not projecting; fore wing shortened, moderately attenuate, exceeding posterior margin of T3; female T1 with moderate hump; T2 with costae not exceeding basal half of tergite; T3 almost entirely smooth; female A1-A4 bright orange-yellow.
Females of H. laticeps can be distinguished from H. parvipennis in having A 1-A4 bright orange-yellow (in H. pawipennis only A1-A2 are bright orange-yellow) and by the upper apices of the propodeal keels being closely approximated (wide apart in H. parvipennis). Male A1-A4 are dark yellow, almost concolourous (male A2 is lighter than A3-A4 in H. pawipennis). Holoteleia laticeps differs from H. polita principally by the entirely sculptured scutellum, by the upper half of the frons being densely coriaceous, and by the presence of a low hump on female TI.
Mesosoma in dorsal view longer than wide (51:40); mesoscutum entirely scalycoriaceous with dense appressed hairs; notaulus fine, deeply incised, almost complete, becoming indistinct at anterior apex; humeral and suprahumeral sulci deep, non-foveolate; transscutal suture with 3 foveolae laterally; scutellar disc entirely coriaceous with scattered setigerous punctures and with dense decumbent hairs; posterior margin of scutellum (in front of scutellar rim) with distinct foveolae; ventral lip of dorsellum smooth, relatively broad; propodeal keels meeting below dorsellum, anterodorsal apices of keels not pointed and not projecting; posterolateral comers of propodeum sharply pointed and projecting; mesosoma in lateral view convex dorsally, longer than high (51:40); side of pronotum mostly smooth with patch of dense microsculpture in front of spiracle and with row of irregular foveolae along dorsal margin; netrion smooth with distinct crenulae along inner margin, crenulae smaller than half netrion width; mesopleural depression smooth and glabrous; mesopleural carina completely smooth, without foveolae ventrally; mesepisternum below mesopleural carina mostly smooth, with scattered fine pilosity and patch of fine coriaceous sculpture below acetabular carina; acetabular carina sharp, not foveolate ventrally; mesepimeron divided from mesepisternum by row of deep foveolae; metapleuron smooth and glabrous with row of foveolae along anterior margin and with deep non-foveolate sulcus; fore wing shortened, moderately attenuate, exceeding posterior margin of T3, with 9 large semierect black bristles on submarginal vein; marginal vein slightly shorter than stigmal vein (6:8),with 2 large bristles, postmarginal vein longer than marginal vein (17:6), with 5 large bristles; median and basal vein not indicated.

Material
Holoteleia parvipennis can be distinguished from the sympatric H. bicolor by the absence of foveolae or irregular sculpture below the mesopleural carina, by the shorter longitudinal costae on T2, and by the absence of coriaceous sculpture on T3. From H. laticeps it differs in the female by having only A1-A5 bright orange-yellow and by having the upper apices of the propodeal keel distinctly apart.
Mesosoma in dorsal view longer than wide (60:46); cervical part of pronotum not foveolate; mesoscutum evenly densely scaly-coriaceous, with dense appressed yellowish hairs; notaulus delicate, incomplete, but well impressed in posterior three-quarters of scutum; humeral and suprahumeral sulci deep, non-foveolate; scutellum anteriorly with 4 deep foveolae and with deep transscutal suture medially; scutellum with even dense coriaceous sculpture, sculpture becoming finer posteriorly, scutellum with scattered decumbent hairs, posterior margin of scutellum (in front of scutellar rim) distinctly foveolate; dorsellum with smooth ventral lip very narrow; propodeum anterodorsally (below dorsellum) deeply excavate with anterodorsal points wide apart, posterior margin of propodeum not carinate, not projecting in sharp points; mesosoma in lateral view convex, longer than high (60:40); side of pronotum predominantly smooth with patch of dense microgranular sculpture in front of spiracle, upper margin of pronotum (below mesoscutum) with row of irregular foveolae, anterior margin of pronotum with band of fine coriaceous sculpture; netrion smooth, foveolate along anterior margin, foveolae smaller than half netrion width; mesopleuron depression smooth and glabrous; mesopleural carina complete, sharp, without foveolae ventrally; mesepistemum below acetabular carina with fine coriaceous sculpture; acetabular carina sharp, not distinctly foveolate ventrally; mesepimeron divided from mesepisterum by row of deep foveolae; metapleuron smooth and glabrous with row of foveolae along anterior margin and with deep non-foveolate median sulcus; fore wing attenuate, reaching only to middle of T3, with 11 semierect bristles on submarginal vein, marginal vein as long as the short and very slanted stigmal vein (6:6); postmarginal vein clearly defined, twice as long as marginal vein (12:6); basal and median veins not indicated.
Metasoma in dorsal view longer than wide (110:41); T1 wider than long (22:17) with strong longitudinal costae except for large smooth area on hump anteromedially with several erect bristles at side; T2 wider than long (38:26), with strong longitudinal costae not reaching basal half of tergite, major part of tergite smooth with few scattered hairs laterally; T3 transverse, wider than long (41:33), smooth, lustrous with only minute patch (only 10 cells) of microcoriaceous sculpture posterolaterally, and with very few scattered hairs around; T4 and T5 smooth with few scattered setigerous punctures, T6 largely smooth with denser pilosity; T7 with pair of bristles on cercus.

Material
Biology. Host unknown. Adult wasps prefer open, sunny habitats such as grassland, sand pits, etc., with peak of occurrence in late summer and early fall.
Variation. Fore wings in females are frequently shortened, not exceeding the middle of T3 but also fully developed; wings are rarely shortened in males. Some male individuals may be dark brown, i.e. much darker than most populations.
Holoteleiapolita is unique among North American species of Holoteleia because of the smooth scutellum and the entirely smooth T3. It differs from H. laticeps which also has the female A1-A4 orange-yellow by the absence of a hump of female T1 and by the frons being predominantly smooth.
Mesosoma in dorsal view longer than wide (5 1 :41); mesoscutum evenly densely scalycoriaceous with dense appressed hairs; notaulus complete, deeply incised particularly in posterior half; humeral and suprahumeral sulci deep, non-foveolate; transscutal suture with 3 4 foveolae laterally; scutellar disc predominantly smooth and glabrous, with only few scattered setigerous punctures and remnants of fine coriaceous sculpture along anterior margin, posterior margin of scutellum (in front of scutellar rim) with distinct foveolae; ventral lip of dorsellum moderately developed; propodeal keels almost meeting below dorsellum, anterodorsal apices of keels not pointed, not projecting; posterolateral comer of propodeurn not pointed; mesosoma in lateral view convex dorsally, longer than high (5 1 :40); side of pronotum largely smooth and glabrous with patch of dense microsculpture in front of spiracle; netrion smooth, foveolate only in lower part, foveolae very small, smaller than half netrion width; mesopleural depression smooth and glabrous; mesopleural carina strong, complete, without foveolae ventrally; mesepistemum below mesopleural carina entirely smooth and glabrous; acetabular carina sharp, not foveolate ventrally; epimeron divided from mesepistemum by row of deep foveolae; metapleuron smooth and glabrous with row of foveolae along anterior margin and with deep non-foveolate median sulcus; wing shortened, attenuate, reaching posterior margin of T3, with 9 large semierect black bristles on submarginal vein; marginal vein longer than short stigma1 vein; postmarginal vein slightly longer than marginal vein; basal and median veins not indicated.
Etymology. Polita is derived from the Latin politus meaning smooth, and referring to the smooth scutellum.