A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)

Systematic Entomology (1989) 14,275-298

A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)

DAVID WAHL American Entomological Institute, Gainesville, Florida

ABSTRACT. The campoplegine genus Benjuminiu is herein revised, with ten species being described as new, in addition to the five previously known species. Phylogenetic relationships of Benjuminia to other genera are discussed, and a cladogram presented for the species. Two monophyletic species groups are recognized, one being confined to the Palearctic and the other to the Nearctic. Examination of larval morphology not only fails to reveal larval characters for distinguishing Benjuminiu but leads to the con- clusion that use of larval morphology to identify campoplegine species is a dubious practice.

Introduction

Benjaminia is a small campoplegine genus restricted to the Holarctic, consisting of fifteen species. Two monophyletic species groups, the fumigator and fuscipennis groups, are recognized. They are confined to the Palearctic and Nearctic, respectively. Benjaminia was erected by Viereck in 1912 for Charops fuscipennis Pro- vancher. Cushman synonymized Zachrestoides Viereck, 192.5, with Benjaminia in 1933. The genus was considered to be Nearctic until publication of Townes’ key to world campoplegine genera (1970) which allowed recognition of the genus by European workers and subsequent description of three Palearctic species (Aubert, 1971; Sawoniewicz, 1973; Kasparyan, 1976).

The biology of Benjaminia is of particular interest. Like all other campoplegines, Ben- jaminia species are koinobiont endoparasitoids. Most campoplegines, while having narrow host ranges, do not confine themselves to a

Correspondence: Dr David Wahl. tornological Institute, 3005 SW 56th ville, FL 32608, U.S.A.

American En- Ave., Gaines-

taxonomic group. All reliable rearing records of Benjaminia have been from Nymphalids of the sub-family Melitaeinae. Parasitism of butterflies is also unusual for campoplegines.

Materials, Methods and Terminology

Specimens examined in this study were bor- rowed from or deposited in the following collections and I am indebted to the following curators:

AEIC: American Entomological Institute: Gainesville, Florida (H. and M. Tow nes) ;

CASC: California Academy of Sciences: San Francisco (N. D. Penny);

CNCI: Canadian National Collections: Ot- tawa, Ontario ( J . R. Barron);

CUIC: Cornell University: Ithaca, New York (E. R. Hoebeke);

DBWC: D. B . Wahl Collection: Gainesville, Florida;

FSAG: Faculte des Sciences Agronomiques de 1’Etat: Gembloux. Belgium (C. Thir- ion);

275

KI I I C - : Klaus Horstmann Collection: Wurz- burg. B . R . D .

1 .A(‘M: 1.0s Angelea County Museum o f Natural History: Los Angeles, Califor- nia (R. R. Snelling):

NESC‘: Nancy E. Stamps Collection: Bing- haniton, New York:

OSUC): Oregon State University: Corvallis ( G . L . Parsons);

RSME: Royal Scottish Museum: Edinburgh ( M . R. Shaw):

UC’DC: R. M. Bohart Museum, University of California, Davis (R. 0. Schuster);

UCRC: University of California. Riverside (J. R. Hall);.

USNM: National Museum of Natural History: Washington, D.C. (E. E. Grissell);

VKGC‘: V . K . Gupta Collection: Gaincsville. Florida:

ZMAS: Zoological Museum. Academy of Sci- ences: Leningrad. U.S.S.R. (D. R . Kasparyan).

‘The morphological terminnlcgy for adults is mostly that of Townes (1969). Epicnerniul cur im is used f o r ‘prepectal carina’, genu for ‘temple‘. itiulur . s p r i ~ ’ e for ‘cheek’. posterior trunsvrrse cnrinu o / r iicsothorucic veri tPr for ‘post pec t al carina‘ . and .s~rpruc/ypeul u r w for ‘face’. Wing vein nomenclature is based upon Ross (1936). M c . s o . v o r i i c c and rrzetusomu are used to refer to the apparent thorax and abdomen. respectively. First / r i c ~ t u . s o m u l segment applies to the second true abdominal segment. Rcference to the oricntation o f the various body parts follows .l‘ou.nes (l9hY) in that the legs arc considered t o he stretched out horizontally at right angles to

Thc terminology for the cephalic sclerites o f the mature larva is that of Finlayson (1975) and Shor-t (1978). except that parietrrl hund is used 101- ‘ocular line’. Methods of preparation are those o f Wahl (1984). My personal notation for larval preparations follows thc museum act-on!m. I t consists o f the day. month. year. and ;I letter designating the individual preparation. f-or example. ‘DBW 17.VII.Xhd’ i s the lourth preparation o f 17 July 1986.

C‘oloi-ation of the body parts. especially that o f thc nietasoma. can be affected by the ;iniwnt o f

the hod!

oils on the specimen (usually from internal sources); o i l y specimcns are usually darker and often exhibit irregular fuscous areas on the nor- mally brownish red portions of the metasoma. Reference to metasomal colour in the descriptions refers only to the tergites and first sternite.

T2, T3. e te . , is used for the second metasomal tergite and following tergites and S2, S3, etc., for the second metasomal sternite and following sternites.

When the lengths of the body and wing are given, the values in parentheses are those of the holotype.

The species descriptions are divided into nine numbered sections for ease of reference. The following are explanations of measurements and characters as needed:

(1) Hypostomul-mandibular inde.u is the length of the hypostomal carina from its juncture with the occipital carina t o the mandibular base, divided by the basal width of the mandible. This is used in place of the ‘length of the genal inflec- tion’ used by Shaw (1977).

(6) Mesopleural punctation is measured in the area immediately below the hypoepimeron (‘speculum‘ of Townes). Whenever surfaces are referred to as smooth, this denotes a complete lack of granulation or other surface sculpture. Size of the mesopleural punctures is in reference to the lateral ocellus diameter (its greatest transverse measurement): fine punctures are 0.1 x lateral ocellar diameter or smaller, mod- erutr punctures are 0.2X lateral ocellar diameter. and lurge punctures are 0 . 3 x lateral ocellar diameter.

Shaw’s (1 977) study o f B. fiimiguror and polonicu used several characters not used here. The length of the intercubitus relative to the length of vein M between the intercubitus and vein 2m-cu in the forewing is subject to significant variation within a species. The same was found to be true for the emargination of the posterior carina of the mesothoracic venter (‘post- pectal carina’). lengthiwidth of the hind femur, and lengthiwidth of T2. While the shapes of the various areas delimited by propodeal carinae are usually quite stable in campoplegines and of great taxonomic use. Renjaminirr is remarkable for the amount of variation in carinae and gen- eral surface sculpture. Although there appears to be significant differences in the overall rugosopunctate sculpture in some species, morc specimens may show that this is unstable as well.

Revision of Benjnminia 277

Taxonomy

Genus Benjaminia Viereck

Benjaminia Viereck, 1912: 633. Type species: Charops fuscipennis Provancher. Original designation.

Zuchrestoides Viereck 1925: 177. Type species: Zachrestoides euphydryadis Viereck. Original designation. Synonymized by Cushman (1933).

Benjarniniu (Fig. 1 ) may be distinguished from other campoplegines by the following: eye weakly indented opposite antenna1 socket; occipital carina joining hypostomal carina above base of mandible; mandible short, ventral margin with narrow lamella which terminates rather abruptly near apex of mandible; ventral tooth of

mandible slightly smaller than dorsal tooth: mesopleurum with fine to large punctures, usually on smooth surface; epicnemial carina somc- times produced ventrally; posterior transvcrsc carina o f mesothoracic venter complete: sublat- era1 carina of propodeum basally elevated as low crest that overhangs propodeal spiraclc: carinae of propodeum usually apically indistinct or absent, surface rugosopunctate; hind basitarsus without special midventral row o f closcly spaced, short hairs; areolet of forewing absent (vein 3r-m absent), vein cu-a distad of composite vein Rs&M by 0.2,X its length or opposite; second abscissa of vein Cu of hindwing basally incomplete, spectral or nebulous (Mason. 1986). composite vein Cu&cu-a vertical t o slightly in- clivous; glymma of TI present and small, sometimes obsolescent; thyridium of T2 large.

278 David Wahl

subcircular or transversely elliptic, position varying from adjacent base of T2 to separated by up to 0.8 x its longitudinal diameter; metasoma weakly laterally compressed; ovipositor about as long as apical metasomal depth.

Biology

All reliable host records of Benjaminia are from rnelitaeine Nymphalidae. The following sum- marizes what is known of host relationships:

R. euphydryadis-Charidryas harrissi (Scudder). C. nycfeis (Doubleday), Euphydryas phaeton (Drury).

B.franklini-unindcntified melitaeine nymphalid.

R.fumigator-Melitaea didyma (Esper). B. fuscipennis-E. chalcedona (Doubleday),

E. editha (Boisduval). B.pellogonia-E.co1on (Edwards), E.editha, E .

editha luesthepa Murphy & Ereich, E.gillettii (Barnes).

B. whifei-E.anicia (Doubleday), E.chalcedona, E.colon, E.editha, E.editha nubigena (Behr).

B.shawi and pellogonia are recorded as being reared from Colius [Pieridae] and Arctiidae, respectively. The reasons why these records are considered dubious are discussed in the treat- ment of each species.

The extent to which a particular Benjaminia species is specific to a host is uncertain. Host specificity is in most cases probably a function of what host i s available and preference for the host o n which the parasitoid developed (Vinson,

What little is known of Benjaminia behaviour is reported in Stamp (1982, 1984). In Virginia, B. euphydryadis attacked E.phaeton early instar larvae in communal webs on Chelone glabra L. [Scrophulariaceae], overwintering as larvae in diapausing fourth instars of phaeton. Fourth instar hosts feed from April to May, then pupate. B.ruphydryadis kills the host just before pupa- tion, the wasps pupating inside the host cuticle for 2-3 weeks before emerging. There is only one generation of euphydryadis per year. B. pellogonia parasitizing E.gillettii in Wyoming (Bear-tooth Mts.) synchronize adult emergence with caterpillars hatching in August. This host population takes two winters to reach maturity. as do the wasps.

1976).

Many species in the Hyposoter complex of genera, to which Benjaminia belongs (see ‘Phylogenetic Relationships’), pupate within the host cuticle. The host remains have an inflated appearance due to the cocoon which is spun within. Benjaminia exhibits the same characters. The cocoon is of an unusually dense and hard texture, unlike any that are known to me. Few loose threads of silk can be found on the exterior or interior surfaces.

Mature larvae of Benjaminia

Mature larval remains of six Benjaminia species (euphydryadis, franklini, fumgator, fuscipennis, pellogonia, whitei) were examined. Short (1978) characterized Benjaminia has having a ‘lightly sclerotized band extending dorsally from each pleurostoma across to meet ventro-lateral edge of antenna’ or ‘postoccipital ridges extending dorsally from posterior tentorial pits to meet in mid-dorsal line.’ The former character, which might be more accurately stated as having an area of variable sclerotization extending dorsally from the anterior tentorial pit to the ventral margin of the antenna, appears to be a synapomorphy of the fuscipennis species group. It is not found in franklini or fumigator. It is somewhat variable intraspecifically. The character of the postoccipital ridge extension is a misinterpreta- tion by Short of fumigator morphology. The original slide mount shows that the dorsolateral extension of the hypostoma (postoccipital ridge of Short) ends about at the level of the antenna1 mid-height; Short apparently thought the parietal band was part of thk hypostoma (Short, 1978). The morphology is similar to other Ben- jaminia larvae. There are no known synapomorphies distinguishing Benjarninia larvae from those of Hyposoter and related genera, which also have the hypostoma dorsolaterally extended and labial lateral areas. Existing keys to campoplegine larvae (Finlayson, 1975; Short, 1978) will not always allow accurate identification of Benjaminia larvae (or those of other genera) and should be used with caution.

Studies of ichneumonid larvae are for the most part typological, based on one specimen which is then used to characterize a species or genus. While this is mostly unavoidable, a cauti- ous attitude is necessary. Much emphasis has

Revision of Benjarninia 279

b -

FIG. 2. Mature larva of B.euphydryadis: (a) cephalic sclerites of 14.11.88~ and d , (b) labial sclerite of 14.11 .XXd. FIG. 3 . Cephalic sclerites of mature larva of B.franklini, lX.II.88b. Scale lines=0.1 mm.

280 David Wahl

FIG. 3 . Cephalic sclerites of mature larva of B.furnigrrror, JRTS n o . 809.

FIG. 5 . Mature larva of B.fuscipennis: (a) cephalic sclerites of 14.11.88b. (b) labial sclerite of 14.11.88a Scale lines=0.1 mm.

Revision of Benjaminia 281

FIG. 6. Mature larva of B.prllogoniu: (a) cephalicscleritesof 16.11.88d. (b) labial scleriteof 16.11.88e. Scale line=0.1 mrn.

been place on various dimensions of the labial sclerite in campoplegines but study of Ben- jarninia reveals this to be a rather plastic structure. While it might be argued that what seems to be a single species based on adult characters actually represents several species with the larvae showing the differences, I consider this very unlikely for euphydryadis and fuscipennis. Un- less one has large series to establish invariant morphologies, use of ratios involving the labial sclerite (and probably other features as well) should be used with caution or avoided.

No reliable larval characters were found to distinguish species except in euphydryadis, where the hypostomal spur is wider and rather blunt at the ventral apical margin.

Comments on each species the sources of specimens are as follows:

euplqdryodis (JRTS no.24; DBW 14. II.88c,

14.II.88d, 16.11.88a; all USNM). Short’s preparation was badly mounted and the cephalic region ripped apart. The specimen’s labial sclerite is similar to that of Fig. 2(a), a composite of 14.11.88~ and 16.11.88a. Short’s figure of the specimen (1978) does not accurately show the hypostomal spur. The associated adult could not be found but the hypostomal spur leaves little doubt as to the proper identification. Fig. 2(a) does not have sclerotization between the anterior tentorial pit and superior mandibular ar- ticulation, but it is present in the other larval preparations of this species.

franklini (DBW 18.11.88b, ZMAS). Shown in Fig. 3.

furnigaror (JRTS no.809, RSME). I com- mented above on Short’s interpretion of the parietal band and dorsolateral extension of the hypostomal carina; see Fig. 4.

282 David Wahl

FIG. 7. Mature larva of R.whitri: (a) cephalic sclerites of 16.11.88h. (h) labial sclerite of 16.11.88~. Scale line=O.l mm

fuscipenriis (DB W 14. II.88a, 14. II.88b, US- NM). The variable relationship between the prelabial sclerite and the interior ventral margin of the labial sclerite is well illustrated here (Fig. 5 ) . Neither of the two published figures of this species (Finlayson, 1975; Short, 1978) actually belong tofusczpmnis. Short’s specimen, which is whitei, is accompanied by a note to the effect that he was aware of the misidentification. Finlayson’s slide, which from the associated adult was determined to be pellogonia, was not examined.

pellogonia (DBW 16.11.88d, 16. II.88e, US- NM). Shown in Fig. 6.

whitei (JRTS no.23; DBW 16.11.88b, 16. 11.88~; all USNM). Short’s slide has the connec- tion between the prelabial sclerite-interior ventral margin of the labial sclerite intermediate t o the two labial sclerites in Fig. 7. The stem of the prelabial sclerite is distinct and a region of sclerotized integument connects it to the labial sclerite.

Phylogenetic relationships

Hyposoter and related genera (Echthronomas, Eriborus, Genotropis, Lemophagus, Prochas, etc.) were chosen as outgroups. Adults share the following synapomorphies: mandible with a ventral flange; small convex clypeus with margin usually narrowly reflexed; short gena; small areola near propodeal base; ovipositor about as long as depth of metasomal apex. In addition, the known mature larvae have the hypostoma extended dorsally as in Figs 2-7. Further resol- ution of the relationships between Benjaminia and these genera is not a t present possible. Hyposoter is almost certainly paraphyletic and a review of the genus on a world-wide basis is necessary to clarify relationships between its component taxa and the genera mentioned above.

Three autapomorphies of Benjaminia can be identified. They are:

(1) Sublateral carina of propodeum basally


elevated as low crest that overhangs propodeal spiracle.

(2) Carinae of propodeum apically weak or absent, surface rugosopunctate.

(3) Areolet of fore wing absent (vein 3r-m absent).

Seven characters were chosen for phylogenetic analysis of the species, polarities being determined by outgroup comparison with the Hyposoter Group. The plesiomorphic state is in parentheses:

(4) Thyridium of T2 adjacent to base of tergite. (Thyridium usually separated by at least 0 . 5 ~ its median diameter.)

(5) T6-7 (and often TS) fuscous. (T6-7 usually brownish red.)

(6) Tarsal claw long, teeth low and not as high as apex of claw. (Claw shorter, some of teeth as high as apex of claw.)

(7) Epicnemial carina of mesopleurum strongly produced ventrally, height at least 0.4x lateral ocellar diameter. (Epicnemial carina height about 0.1 X

lateral ocellar diameter.) (8) Mature larva with sclerotized area ex-

tending dorsally from anterior tentorial

8-9

Fuscipennis group

pit to ventral margin of antenna. (Sclerotized region absent.)

(9) Supraclypeal area rugulosopunctate. (Supraclypeal area either punctate on smooth surface or granulosopunctate.)

(10) Tegula brownish red or brown. (Tegula usually white.)

Some ambiguities exist regarding characters 4, S and 10. Some Hyposoter species (mostly in the Palearctic) exhibit the apomorphic states of these characters, although none possess two or more simultaneously. Until more is known about Benjarninia’s relationship to other taxa of the Hyposoter Group, I have chosen to regard the more widely distributed character as plesiomorphic. I am mindful of the pitfalls of this approach.

The cladogram that resulted from hand- analysis of the data is shown in Fig. 8. Two sister- groups of species are resolved. The fumigator species group (aridens, franklini, fumigator, horstmanni, kasparyani, maurus, polonica, shawi) is delimited by possession of synapomorphies 4-6. A subgroup consisting of fumigator, maurus, polonica and shawi is delimited by synapomorphy 7. All species in the fumigator

Z - r m i g a t o r group

FIG. 8. Cladogram of Benjarninia species. Double line indicates parallelism.

284 David Wahl

group arc confined to the Palearctic. The Nearc- tic Benjaminia species form a monophyletic group, the fumipennis species group (carlsoni, euphydryadis, fuscipennis, paeminosa, pel- logoniu, phaeothrix, whitei). With the exception of the clade fumigator+maurus+polonica+ shawi. relationships within each species group are unresolved. No doubt this could be improved with more definite knowledge of an outgroup for the genus. It should be pointed out that even if characters 4,s and 10 are found to be incorrecctly polarized, monophyly of the species groups is still upheld.

Key to species of Benjaminia

Tarsal claw long, teeth low and not as high as apex of claw (Fig. 10); T5-8 fuscous; thyridium adjacent to base of T2; tegula almost always white: Paearctic. FUMIGATOR G R O U P ............ 2 Tarsal claw shorter, some of teeth as high as apex of claw (Fig. 9); T2-8 brownish red; thyridium separated from base of T2 by 0.4-0.8X its diameter; tegula fuscous or brownish red; Nearctic. FUSCIPENNIS G R O U P ......................... 9

Tcgula fuscous; setae of head and mesosoma dark

Tegula white; setae of head and mesosocutum white ................................................... 3 Epicnemial carina produced ventrally, height 0 . 4 ~ lateral ocellus diameter or greater ....... 4 Epicnemial carina not produced, height 0.2X lateral ocellus diameter or less ...................... 7

Hypostomal-mandibular index= 0.7 .... fumigator Hypostomal-mandibular index= 1-1.2 . . . . . . . . . 5

brown ..................................... kasparyanc

Fore coxa predominately brownish red; first lateral area coarsely punctate; wings brown; punctures of hind femur separated by 2-3xtheir diameter ....................................... maurus

Fore coxa fuscous; first lateral area rugosopunctate; wings clear to having faint brown tint; punctures of hind femur separated by 1-3xtheir diameter ............................................... 6 Punctures of mesopleurum moderate to large, subadjacent t o separated by about 0.5X their diameter; basal transverse carina and basal sections of median longitudinal carinae high and strong; wings with faint brown tint, veins dark brown; punctures of hind femur separated by 1-2xtheir diameter ..................................... polonica Punctures of mesopleurum moderate, separated by 0.5-1 .Ox their diameter; basal transverse carina and basal sections of median longitudinal carinae low and rather delicate; wings clear. veins light brown: punctures of hind femur separated by about 3 X their diameter ..................... shawi

11

12

7 Punctures of mesopleurum. moderate to large, confluent to separated by about 0.3x their diameter; scape brownish red .................. franklini Punctures of mesopleurum moderate, separated by 0.5-1 .OX their diameter; scape brownish red or fuscous ................................................. 8

8 Scape brownish red; punctures of mesopleurum separated by about their diameter; punctures of

..................................................... aridens Scape fuscous; punctures of mesopleurum separated by 0.5-1 .Ox their diameter; punctures of

................................................ horstrnanni 9 Setae of head and mesosoma dark brown ._ . 10

Setae of head and mesosoma white ........... 11 10 Second trochanters, femora, and tibiae light brow-

nish red; punctures of mesopleurum moderate,

Second trochanters, femora, and tibiae dark brown: punctures of mesopleurum moderate to large, subadjacent t o separated by 0.2X their

hind femur separated by 1-2x their diameter ......

hind femur separated by 2-3x their diameter ......

separated by 0 . 3 - 0 . 5 ~ their diameter . . . carlsoni

diameter, surface rugosopunctate .... phaeorhrix

Hypostomal-mandibular index = 0.4-0.5; legs brownish red ........................................ 12

Hypostomal-mandibular index= 1 .O-1.3; legs with at least coxae and first trochanter fuscous .. 13

Punctures of mesopleurum fine to moderate, usually separated by 1-2x their diameter (in males, often 0.3-l.0x); wings brown; gonoforceps brow-

Punctures of mesopleurum moderate t o large, suh- adjacent, surface rugosopunctate; wings with

nish red .................................... fuscipennis

brown tint; gonoforceps brown ...... paeminosa

a- - 9 10

11 12 FIG. 9. Tarsal claw of B.fuscipennis. FIG. 10. Tarsal claw of B.arridens. FIG. 1 1. Dorsal aspect of head of B.fumigator. FIG. 12. Dorsal aspect of head of B.po/onica.


13 Femora dark brown-fuscous; clypeus granulosopunctate; punctures of mesopleurum subadjacent, surface rugosopunctate . . . , , , . , , . . . . euphydryadis

Femora brownish red; clypeus with surface between punctures smooth; punctures of mesopleurum separated by 0.3-1.0X their diameter . . 14

14 Apicolateral corners of T2 with transverse fuscous marks; postpetiole usually with at least basal 0.3 fuscous; S8 of male as in Figs 22-24 . . . . pellogonia

Apicolateral corners of "'2 without fuscous marks; postpetiole brownish red; S8 of male as in Figs 25- 27 .................................................. whitei

. .

Species descriptions

1. The fumigator species groups

Benjaminia aridens Kasparyan (Fig. 10)

Benjaminia aridens Kasparyan, 1976: 71. Type:

Benjuminiu aridens Kasparyan, 1981: 420 (key).

DIAGNOSIS. Distinguished from other species in the fumigator group by: hypostomal- mandibular index= 1 .O; scape brownish red; epicnemial carina not produced laterally; setae of head and mesosoma white; tegula white; wings clear; punctures of mesopleurum moderate and separated by about their diameter; punctures of hind femur separated by I-2x their diameter.

FEMALE. Structure. 1. Hypostomal-mandibular index=1.0. 2. Antenna with 32 flagellomeres. 3. Supraclypeal area with punctures confluent to separated by about 0 . 3 ~ their diameter, not appearing rugosopunctate; punctures of clypeus sparse apically, basally separated by about their diameter. 4. Gena in dorsal view strongly receding and weakly convex, similar to Fig. 11. 5. Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum fine to moderate, separated by about their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 1-2x their diameter. 8. First lateral area weakly rugosopunctate, rest of propodeal surface more strongly so; basal transverse carina, and sections of median longitudinal carinae forming basal area and areola, present and strong, remainder of carinae indistinct o r absent.

Colour. Head and mesosoma black, the following brownish red: scape and pedicel, apex of

d[ZMAS]; examined in 1987.

mandible, fore leg, basal 0.5 of middle coxa and remainder of middle leg, hing leg beyond second trochanter. Remainder of middle coxa, hind coxa, and hind first trochanter, piceous. Palpi brownish red. Setae of head and mesosoma white. Tegula white. Wings clear. Basal 0.5 of petiole fuscous; remainder of metasoma brownish red except for dark brown of TS-7.

Length. 9.2 mm; forewing 6.0 mm. MALE. Structure. Similar to female except

clypeus evenly punctate, punctures separated by 0 . 5 - 1 . 0 ~ their diameter; punctures of mesopleurum separated by 0.5-1.0~ their diameter; first lateral area with rugae more pronounced, but not as developed as on rest of propodeum. Colour. As in female except pedicel brown; fore coxa and all trochanters dark brown-fuscous; hind leg with coxa black, basal 0.5 of tibia, and tarsus, dark brown; petiole, basal 0.5 of postpetiole, and gonoforceps fuscous. Length. 8.1 mm; forewing 5.9 mm.

SPECIMENS EXAMINED. Type material. holotype 0, U.S.S.R.: Kazakh S.S.R. , Peski Muyunkum, 27.vii. 1931 (Vel'tischev) (ZMAS). Condition of type: intact. Other specimens. Ar- menia S.S.R., Yerevan, near Berdadzor River, 14.vii.1969 (Rikhter) (18, ZMAS).

COMMENT. T2-4 have irregular darker areas, a preservational artefact

Benjaminia franklini sp.n. (Fig. 3)

DIAGNOSIS. This species can be distinguished from others in the fumigator group by: hypostomal-mandibular index = 1 .O; brownish red scape; epicnemial carina not produced ventrally; setae of head and mesosoma white; tegula white; wings clear; punctures of mesopleurum moderate to large, confluent to separated by 0.3-0.SX their diameter, surface rugosopunctate; punctures of hind femur separated by 1-2X their diameter.

MALE. Structure. 1. Hypostomal-mandibular index = 1.0. 2. Antenna with 32 flagellomeres. 3. Supraclypeal area with punctures confluent to separated by about 0.3x their diameter, not appearing rugosopunctate; punctures of clypeus separated by about 0.5x their diameter. 4. Gena in dorsal view strongly receding and weakly convex, similar to Fig. 11. 5. Epi- cnemial carina not produced ventrally. 6. Punc- tures of mesopleurum moderate to large.

286 David Wahl

confluent to separated by 0.3-0.5 X their diameter. surface rugosopunctate. 7. Punctures of anterior face of hind femur separated by about 1-2x their diameter. 8. Propodeal surface coarsely rugosopunctate; basal transverse carina and portions of median longitudinal carinae forming basal area present and strong, remainder of carinae indistinct or absent.

Colour. Head and mesosoma black, the following brownish red: scape and pedicel, mandible except for brown base, palpi, fore and middle legs. hind legs beyond second trochanter except for brown apical 0.3 of tibia; hind coxa and first trochanter deep brownish red with fuscous patches; propodeum with 2 spots of deep brownish red on second lateral areas. Tegula white. Setae of head and mesosoma white. Wing clear. First metasomal segment fuscous except for brownish red of postpetiole; remainder of metasoma brownish red except for fuscous T5-8 and gonoforceps.

Length. 8.2 mm: forewing 6.0 mm. TYPE MATERIAL. Holotype 8, U.S.S.R.:

Armenia S.S.R., Yerevan, 26. viii.1969 (Ertevt- syan) (ZMAS). Condition of type: intact.

COMMENT. As with some other specimens, it is difficult to tell how much of the dark coloration of the metasomal apex is natural and how much is due to preservational vagaries.

The remains of the host, a melitaeine nymphalid, are pinned under the specimen.

ETYMOLOGY, Named after Benjamin Franklin. American philosopher and statesman.

Benjaminia fumigator Aubert (Figs 4, 12)

Benjaminia fumigator Aubert, 1971: 42. Type:

Benjaminia fumigator Sawoniewicz, 1973: 667-

Benjaminia fumigator Kasparyan, 1976: 70

Benjaminia fumigator Shaw, 1977: 85-89. Host:

Benjarninia fumigator Short, 1978: 80, 351

Benjaminia fumigator Kasparyan, 1981: 420

0 [FSAG]; examined in 1987.

669 (taxonomy).

(key 1.

Melitaea didyma (taxonomy, biology).

(larva).

(key).

DIAGNOSIS. Distinguished from other species in the fumigator group by: hypostomal-

mandibular index=0.7; epicnemial carina ventrally produced; punctures of mesopleurum moderate, separated by 0.3-1.OX their diameter; fore coxae fuscous; wings with brown tint; first lateral area coarsely rugosopunctate; punctures of hind femur separated by about 2~ their diameter.

FEMALE. Structure. 1. Hypostomal-mandibular index = 0.7. 2. (Antennae incomplete). 3. Supraclypeal area with punctures subadjacent, surface not rugulosopunctate; clypeus with punctures separated by about 0.5-1.Ox their diameter, on smooth surface. 4. Gena in dorsal view strongly receding, weakly convex as in Fig. 11. 5. Epicnemial carina strongly produced ventrally, height about 0 . 4 ~ lateral ocellar diameter. 6. Punctures of mesopleurum moderate, subadjacent to separated by about their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by about 2 x their diameter. 8. Surface coarsely rugosopunctate; basal transverse carina and sections of median longitudinal carinae forming basal area and areola present, remainder of carinae indistict or absent.

Colour. Head and mesosoma black; legs beyond second trochanters brownish red except for brown of extreme base of hind femur, apical 0.2 of hind tibia, and hind tarsus; first trochanters with varying amounts of brownish red. Apex of mandible, and palpi, brown. Setae of head and mesosoma white. Tegula white. Wings with brown tint. Petiole fuscous; remainder of metasoma brownish red except for fuscous of TS-7.

Length. 8.8 mrn; forewing 6.3 rnrn. MALE. Structure. As in female, 32 flagello-

meres; some punctures of mesopleurum large, punctures separated by 0.5-1 .OX their diameter; propodeal surface with rugae finer and more numerous, carinae weaker and more irregular. Colour. A s in female, except hind tibia with apical 0.3 dark brown and TS with median brownish red area (T5 fuscous areas irregular, colour may be preservational artefact). Length. 8.6 mm; forewing 6.3 mm.

SPECIMENS EXAMINED. Type material. holotype P , FRANCE: Hautes Alpes, Ville Vieille, 1440 m, 4.viii.1966 (Leclercq) (FSAG). Condition of type: antennae incomplete; dor- sum of mesosoma split open by pin. Other specimens. FRANCE: Var, Draguignan, ex Melitaea didyma, host larva collected 26.v.1974, adult


brown. Tegula white. Setae of head and mesosoma white. Wings with brown tint. First metasomal segment fuscous except for brownish red apical 0.5 of postpetiole; remainder of metasoma brownish red except for fuscous

Length. 8.7 mm (9.7 mm); forewing 6.6 mm (6.4 mm).

MALE. Structure. As in female, antenna with 32 flagellomeres. Colour. Similar to female, hind femur dark brown except for deep brownish red of apical 0.3. Length. 9.2 mm; forewing 6.3 mm.

TYPE MATERIAL. Holotype 9, TURKEY: 10 km S. Ankara, 1050 m (Warncke) (KHIC). Condition of type: intact except antennae incomplete. Paratypes: U.S.S.R.: Kalmyk (??- 200 km E-SE of Voronezh, Russian S.F.S.R.), 31.v.1907 (Borodin) ( 1 9 , ZMAS); Kazakh S.S.R., Janvarzevo, near Ural River, 17.vi.1949 (Rudolf) (1 6 , ZMAS) .

ETYMOLOGY. Named for D r Klaus Horstmann in recognition of his contributions to ichneumonid systematics.

T5-7.

emerged 19.vi.1974 (Shaw) (1 8, RSME); Vauc- luse, near Bedoin, 8.vii.1987 (Graham) ( 1 9 , BMNH).

COMMENT. Although there are some differences in the nature of the mesopleural punctation and propodeal rugosity, the hypostomal- mandibular index, similar colour of the wings, and distinct propodeal rugosity make the assign- ment of the male to fumigator fairly certain. Shaw (1977) remarked on the uniform wing colour of both sexes, as contrasted to the dimorph- ism reported by Cushman (1933) for fuscipennis. Cushman’s finding was in error, as he had mis- taken a male of whitei forfuscipennis.

The Draguignan specimen is the source of the mature larvae figured by Short (1978).

Benjaminia horstmanni sp.n.

DIAGNOSIS. This species can be separated from other species in the fumigator group by: hypostomal-mandibular index= 1 .O; scape fuscous; epicnemial carina not ventrally produced; setae of head and mesosoma white; tegula white; wings with faint brown tint; punctures of mesopleurum moderate and separated by 0.5-1.OX their diameter; punctures of hind femur separated by 2-3x their diameter.

FEMALE. Structure. 1. Hypstomal-mandibular index = 1.0. 2. Antenna with 30 flagellomeres. 3. Supraclypeal area with punctures separated by about 0.3x their diameter; punctures of clypeus separated by 0 .5-1.0~ their diameter. 4. Gena in dorsal view strongly receding and weakly convex, similar to Fig. 11. 5 . Epicnemial carina not produced ventrally. 6. Punctures of mesosoma moderate, separated by 0 . 3 - 0 . 5 ~ their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 2-3x their diameter. 8. First lateral area with punctures subadjacent and only weakly rugose, remainder of propodeal surface coarsely rugosopunctate; basal transverse carina and sections of median longitudinal carinae forming basal area and areola present and strong, remainder of carinae indistinct or absent.

Colour. Head and mesosoma black, the following brownish red: apex of mandible; anterior basal 0.3 of fore coxa; remainder of fore leg after second trochanter, middle leg beyond second trochanter, hind femur except for fuscous basal 0.3. Palpi brown. Hind tibia and tarsus dark

Benjaminia kasparyani sp.n.

DIAGNOSIS. Distinguished from other species in the fumigator group by: hypostomal- mandibular index= 1.0; fuscous scape; epicnemial carina not produced ventrally; setae of head and mesosoma dark brown; fuscous tegula; brown wings; punctures of mesopleurum moderate, separated by about their diameter; punctures of hind femur separated by 2-3 x their diameter.

FEMALE. Structure. 1. Hypostomal-rnan- dibular index=l.O. 2. Antenna with 31-32 flagellomeres. 3. Supraclypeal area with punctures separated by 0.3-1.Ox their diameter or greater; clypeus with same range of punctation, on smooth surface. 4. Gena in dorsal view strongly receding and weakly convex, similar to Fig. 11. 5. Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum moderate, separated by about their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 2-3x their diameter. 8. Propodeal surface coarsely rugosopunctate; basal transverse carinae and sections of median longitudinal carinae forming basal area and areola present and strong, remainder of carinae indistinct o r absent.

288 David U'ahl

Colour. Head and mesosoma black; apex of mandible and legs beyond second trochanter, with exception of dark brown hind tibia and tarsus, brownish red. Tegula fuscous. Setae of head and mesosoma dark brown. Wings brown. First metasomal segment fuscous except for brownish red of apical 0.2-0.3 of postpetiole; T2-4 brownish red, T5-7 and gonoforceps fuscous.

Length. 8.2-9.0 mm (9.0 mm); forewing 6.0- 6 .2mm (6.2mm).

TYPE MATERIAL. Holotype d, U.S.S.R.: Tadzhik S.S.R., Kondara (asmall reserve), near Varzob River, 1100 m , 17.vi.1937 (Cus- sakovskij) (ZMAS). Condition of type: intact. Paratypes, U.S.S.R.: Tadzhik S.S.R., 30 km NW Ramit, 10-12.vi.1975 (Pesenko) ( 1 6 , ZMAS); Tadzhik S.S.R., Kondara, 19.v.1939 (Gussakovskij) (1 d , AEIC).

ETYMOLOGY. Named for D r D. R. Kaspa- ryan in recognition of his contribution to ichneumonid systematics.

Benjaminia maurus spn.

DIAGNOSIS. Distinguished from other species in the fumigator group by: hypostomal- mandibular index= 1 .O; epicnemial carina ventrally produced; punctures of mesopleurum separated moderate and separated by about their diameter; wings brown; fore coxae brownish red; first lateral area coarsely punctate; punctures of hind femur separated by 2-3x their diameter.

FEMALE. Structure. 1 . Hypostomal-mandibular indcx=l .0 . 2. Antenna with 32 flagellomeres. 3. Supraclypeal area with punctures subadjacent to separated by about 0.5X their diameter; clypeus with punctures separated by about their diameter. 4. Gena in dorsal view weakly receding and not convex, similar to Fig. 12. 5 . Epicnemial carina strongly produced ventrally, height about 0.4X lateral ocellar diameter. 6. Punctures of mesopleurum moderate and separated by about their diameter, on a smooth surface. 7. Punctures of anterior face of hind femur separated by 2-3 X their diameter. 8. First lateral area with large punctures and no traces of rugae. remainder of propodeal surface rugosopunctate; basal transverse carina and sections of mcdian longitudinal carina forming basal area and areola present, remainder of carinae weak or abscnt .

Colour. Head and mesosoma black, the following brownish red: fore reg except for extreme base of coxa and basal 0.5 of first trochanter, middle femur and tibia, hind femur and tibia except for brown of apical 0.4 of tibia; apex of middle coxa, middle and hind trochanters, with indistinct brownish red areas. Palpi brown. Setae of head and mesosoma white. Tegula white. Wings brown. Petiole fuscous; remainder of metasoma brownish red except for fuscous of T6-7 (all tarsi missing).

Length. 9.9 mm; forewing 6.9 mm. TYPE MATERIAL. Holotype 0 , MOR-

OCCO: 'Daiet Aoua', lS.vi.1961 (AEIC). Con- dition of type: right antenna incomplete; right fore leg beyound coxa missing; all tarsi missing; posterior left hind tibia1 spurs missing.

COMMENT. H. Townes (pers. comm.) does not remember the provenance of this specimen. The location cannot be found in standard at- lases, but is perhaps Dayat et Aouda [32"47'N, 7" 2O'W] (Morocco Oficial Standard Names Gazetter, U.S. Board in Geographic Names).

ETYMOLOGY. From the Latin Maurus, a native of North Africa or Moor. It is a noun in apposition.

Benjaminia polonica Sawoniewicz (Fig. 12)

Benjaminia polonica Sawoniewicz, 1973: 667- 669. Type: 9 Museum of the Institute of Zoology, Polish Academy of Sciences, Warszawa]; type not examined.

Benjaminiupolonica Kasparyan, 1976: 70 (key). Benjaminiu polonica Shaw, 1977: 85-89

Benjaminia polonica Kasparyan, 1981 : 420 (taxonomy).

(key).

DIAGNOSIS. This species can be separated from others in the fumigator group by: hypostomal-mandibular index= 1.2; epicnemial canna ventrally produced; punctures of mesopleurum moderate to large, subadjacent to separated by 0.Sx their diameter; fore coxa fuscous; wings with brown tint; punctures of hind femur separated by l-2x their diameter; first lateral area coarsely rugosopunctate.

FEMALE. Structure. 1 , Hypostomal-mandibular index=1.2. 2. Antenna with 32-33 flagellomeres. 3. Supraclypeal area with punctures subadjacent, surface not ruguloso-


punctate; clypeus with punctures separated by about 0.5-1.OX their diameter, on smooth surface. 4. Gena in dorsal view weakly receding, not convex (Fig. 12). 5. Epicnemial carina strongly produced ventrally, height about 0.6X lateral ocellar diameter. 6. Punctures of mesopleurum moderate to large, subadjacent to separated by 0 . 5 ~ their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 1-2X their diameter. 8. Surface coarsely rugosopunctate; basal transverse carina and sections of median longitudinal carinae forming basal area and areola present, remainder of carinae indistinct o r absent.

Colour. As for fumigator Aubert, except: brownish red of legs and metasoma deeper in tint; apical 0.3 of hind tibia brown; wings with very faint brown tint, veins light brown; first metasomal segment fuscous except for brownish red apical 0.5 of postpetiole.

Length. 8.5-8.8 mm; forewing 6.2-6.3 mm. SPECIMENS EXAMINED. Paratype 9,

POLAND: Bilaystok-Pietrasze, on flowers of Peucedonum oreoselinum, 4.viii. 1969 (Sawo- niewicz) (AEIC); ‘Germany; Ruthe Coll.; 59.101’ ( 9 , BMNH).

COMMENT. The type series was collected on Peucedonum in 2-3-year cultures of Scots pine in a mixed coniferous forest (Sawoniewicz, 1973).

Benjaminia shawi sp.n.

DIAGNOSIS. This species can be distinguished from others in the fumigator group by: hypostomal-mandibular index= 1.0; epicnemial carina produced ventrally; punctures of mesopleurum moderate, separated by 0.5-1.OX their diameter; fore coxae fuscous; wings clear, veins light brown; first lateral area delicately rugosopunctate; punctures of hind femur separated by about 3x their diameter.

FEMALE. Structure. 1. Hypostomal-mandibular index=l.0. 2. Antenna with 31 flagellomeres. 3. Supraclypeal area with punctures separated by about 0 . 5 ~ their diameter. 4. Gena in dorsal view strongly receding and weakly convex, similar to Fig. 11. 5 . Epicnemial carina strongly produced ventrally, height about 0.5 X

lateral ocellus diameter, on smooth surface. 7.

Punctures of anterior face of hind femur separated by 0.3-0.5x their diameter. 8. Propodeal surface rugosopunctate, rugae low and delicate; basal transverse carina and sections of median longitudinal carinae forming basal area and areola present but low and delicate, remainder of carinae indistinct or absent.

Colour. Head and mesosoma black, the following brownish red: apex of mandible, fore leg beyond first trochanter, middle and hind legs beyond second trochanter except for brown of apical 0.2 of hind tibia, and hind basitarsus (hind tarsi incomplete); middle and hind second trochanters with varying amounts of brownish red. Palpi brown. Setae of head and mesosoma brownish red. Tegula white. Wings clear, veins light brown. Petiole fuscous; remainder of metasoma brownish red except for fuscous of T5-7 (T5 with median brownish red area, fuscous coloration may be preservational artefact.

Length. 7.3 mm; forewing 6.7 mm. TYPE MATERIAL. Holotype ? , FRANCE:

Vaucluse, Mt Ventoux, vi.1974 (Charmon & McLeod) (RSME). Condition of type: right antenna missing except for pedicel and scape, left antenna mounted on point below specimen; tarsomeres 4-5 of left fore leg missing; right fore- and hindwings detached and point below specimen; tarsomeres 2-5 of right and left hind leg missing.

COMMENT. The adult and putative host pupa were said to be reared from a pupa of Col- ias croceus (although the associated-sent later (M. R. Shaw, pers. comm.)-was labeled C.au- stralis. I examined the Colias chrysalid for Ben- jaminia larval remains. Inside were: (1) a large white meconial mass unlike those found with other Benjaminia larvae; (2) remains of a pupa that appears to be that of a large chalcidoid (H. Townes, pers. comm.); (3) cast larval skin of a parasitoid Hymenoptera, as large as a mature Benjaminia cast larval skin but lacking the usual sclerotized features of ichenumonoid larvae-the only prominent features were the large mandibles. These observations, along the with host being a pupa and not a larva, would indicate that some sort of mistake was made in associating host remains with the Benjaminia adult. Perhaps the wasp did emerge from a Colias but the wrong pupa was chosen.

ETYMOLOGY. Named after Dr M. R. Shaw for his work in parasitoid Hymenoptera biology.

290 David Wahl

2. The fuscipennis species group

Benjaminia carlsoni sp.n. (Fig. 27)

DIAGNOSIS. Distinguished from other species in the fuscipennis group by: setae of head and mesosoma dark brown; hypostomal-mandibular index=0.6; wings with brown tint; legs beyond first trochanters brownish red; punctures of hind femur separated by 3-5X their diameters.

FEMALE. Structure. 1. Hypostomal-mandibular index=0.6. 2. Antenna with 35 flagellomeres. 3. Supraclypeal area rugosopunctate; clypeus with punctures separated by about 0.5 X

their diameter on smooth surface. 4. Gena in dorsal view moderately receding and weakly convex, similar to Fig. 17. 5. Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum moderate, separated by 0.3-0.5x their diameter (ocasionally 0.8-1.0X), on smooth surface. 7. Punctures of anterior face of hind femur separated by 3 - 5 ~ their diameter. 8. Sur- face of propodeum finely rugosopunctate; carinae present except for median longitudinal carinae apicad, the basal transverse carina.

Colour. Head and mesosoma black, the following brownish red: palpi and legs beyond first trochanters, with exception of brown hind tarsus. First trochanters fuscous except for brownish red of extreme apices. Tegula dark brown. Wings with brown tint. Setae of head and mesosoma dark brown. Basal 0.8 of first metasomal segment fuscous, remainder of metasoma brownish red.

Length. 11.1 mm; forewing7.1 mm. TYPE MATERIAL. Holotype 9 , UNITED

STATES: Oregon, Jackson Co. , Pinehurst (5 miles S. Hyatt Reservoir on Route 66) , 1Y.vi.1978 (Townes) (AEIC). Conditionof type: intact.

ETYMOLOGY. Named for Dr R. W. Carlson in recognition of his preliminary taxonomic work on the genus

Benjaminia euphydryadis (Viereck) (Figs 2, 13. 15, 19)

Zmhwstoides euphydryadis Viereck, 1925. 1926: 179 (key); 58: 3. Type: P (CNCI); examined in 1987. Host: Euphydryusphaeton.

Benjarninia euphydryadis Cushman, 1933: 16 (taxonomy, biology)

Benjaminia euphydryadis Schaffner & Griswold, 1934: 152. Host: E.phaeton, Charidryas har- risii (biology).

Benjaminia euphydryadis Short, 1978: 80, 352 (larva).

Benjaminia euphydryadis Stamp, 1982: 100 (biology.

Benjaminia euphydryadis Stamp, 1984: 2-18 (biology).

DIAGNOSIS. Distinguished from other species in the fuscipennis group by: setae of head and mesosoma white; hypostomal-mandibular index=l . l ; surface of clypeus and mesothoracic venter granulate; mesopleurum rugosopunctate; wings clear; middle and hind femora brown-fuscous; punctures of hind femur separated by 1-2X their diameter.

FEMALE. Structure. 1. Hypostomal-mandibular index=l . l . 2. Antenna with 34-35 flagellomeres. 3. Supraclypeal area centrally granulosopunctate, punctures adjacent to separated by about 0.3 X their diameter; punctures of clypeus separated by 0.3-0.5 X their diameter on granulate surface. 4. Gena in dorsal view straight and only slightly receding (Fig. 15). 5 . Epicnemial carina not ventrally produced. 6. Punctures of mesopleurum fine, separated by 0.5-1.Ox their diameter, on granulate surface, surface appearing rugosopunctate. 7. Punctures of anterior face of hind femur separated by 1-2x their diameter. 8. Surface of propodeum rather finely rugosopunctate; basal transverse carina and sections of median longitudinal carina forming basal area and areola present, remaining carinae indistinct or absent.

Colour. Head and mesosoma black, with dorsal and posterior faces of fore and middle femora, and hind tibia except for narrow apical fuscous stripe on anterior and posterior faces, deep brownish red. Fore first trochanter, dorsal surface of fore femur, fore tibia, apical 0.1 of middle femur, and basal 0.2 of middle tibia, yellowish white. Fore and middle tarsi except for apical tarsomeres and remainder of middle tibia, brownish white. Remaining trochanters fuscous. Middle and hind femora, with above exceptions, dark brown-fuscous. Tegula brown with whitish apex. Wings clear. First metasomal segment with petiole and basal 0.5-0.8 of postpetiole fuscous; remainder of metasoma brownish red.


Length. 6.0-10.3 mm (6.0mm); forewing4.2- 7.9mrn(4.2 mm).

MALE. Similar to female except antenna with 33-34 flagellomeres; surface sculpture and carinae of propodeum somewhat more developed; S8 as shown in Fig. 19. Colour. As in female, except yellowish white areas of legs replaced by brownish white; brownish red areas of middle and hind legs replaced by dark brown- fuscous; first metasomal segment completely fuscous; basal 0.2 of T2 and gonoforceps, fuscous. Length. 9.20-10.7 mm; forewing 6.5-7.2 mm.

SPECIMENS EXAMINED. Type material. Holotype 9 , CANADA: Quebec, St. Adele, ex Euphydryas phaeton (CNCI, type no. 1579). Condition: intact except left hind tarsus with tarsomeres 4-5 missing, Other specimens. CANADA: Ontario, Stittsville, 2l.vii. 1975 (Sunborne) (1 0 , AEIC); UNITED STATES: Massachusetts, Middlesex Co., Reading, ex Charidryas nycteis, ‘GipMothLab 12411E2’, 7.vii.1919 ( Id , USNM]; Essex Co., Saugus, ex E.phaeton, ‘GipMothLab 12410N2’, (cocoon attached to separate pin), 3.viii.1927 (1 9, USNM); same data as above but ‘GipMothLab 12410S1 and collected 26.vi.1930 (1 P , USNM); New York, Tompkins Co., Ithaca, Cornell Res.

Park, ex Ephaeton, vii.1978 (White) (19, 1 8 , USNM); Tompkins Co., Ithaca, (Smith) (1 sex?, CUIC); Virginia, Warren Co., Front Royal, ex Euphdryas phaeton larva, 28.v.1979, l.vi.1979, 3.vi.1979 (19, 3 8 , NSIC).

COMMENT. Females can have leg coloration varying from what is described above t o a dark condition as found in males. A male from Front Royal, Virginia, has the supraclypeal area and clypeus not distinctly granulate, the punctures appearing to be obliterated.

Benjaminia fuscipennis (Provancher) (Figs 1 , 5 , 9 , 14, 16,20)

Charops fuscipennis Provancher, 1888: 365. Type: 9 (USNM); examined in 1987 Host: Euphydryadis chalcedona.

Benjaminia fuscipennis Viereck, 1912: 633 (taxonomy).

Benjaminia fuscipennis Gahan & Rowher, 1917: 336 (taxonomy).

Benjaminia fuscipennis Cushman, 1933: 16 (taxonomy, biology).

Benjaminia fuscipennis Barron, 1975: 475 (taxonomy).

~~ ~

FIG. 13. Known distribution of B.euphydryadis. FIG. 14. Known distribution of B.fuscipennis.

292 David Wahl

DIAGNOSIS. This species can be distinguished from others in thefuscipennis group by: setae of head and mesosoma white; hypostomal- mandibular index=0.4; punctures of mesopleurum tine to moderate, separated by 1-2x their diameter in females; wings brown; tegula and legs beyond coxae brownish red; punctures of hind femur separated by 4-5 x their diameter; gonoforceps brownish red.

FEMALE. Structure. 1. Hypostomal-mandibular index=0.4. 2. Antenna with 35 flagellomeres. 3. Supraclypeal area centrally with fine punctures, separated by about 1 - 2 ~ their diameter, laterally rugulosopunctate; punctures of clypeus separated by 0.5-1 .OX their diameter, on smooth surface. 4. Gena in dorsal view strongly convex and sharply receding posteriorly (Fig. 16). 5 . Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum fine and separated by 1-2x their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 4-5x their diameter. 8. Sur- face of propodeum coarsely rugosopunctate; propodeal carinae present, varying from complete and strong, to almost indistinguishable from surface sculpture.

Colour. Head and mesosoma black, the following brownish red: ventral surfaces of scape and pedicel, mandibles except for fuscous of extreme base and apex, palpi, tegula, apical 0.5 of fore coxa. apical 0.3 of middle coxa, extreme apex of hind coxa, remainder of legs. Wings brown. Setae of head and mesosoma white. Metasorna with basal 0.5-0.7 of petiole piceous- fuscous; remainder of metasoma brown.

Length. 9.8-12.2 rnm(9.8mm); forewing7.1- 8 . 4 m m ( 7 . 2 mm).

MALE. Structure. Similar to female except antenna with 35-38 flagellomeres; central region of supraclypeal area rugulosopunctate; mesopleural punctures moderate and separated by 0.5-1 .OX their diameter; S8 as shown in Fig. 20. Colour. Similar to female except gonoforceps brownish red. Length. 9.0-10.1 mm; forewing 6.6-7.9 mm.

TRIBUTION, Type material. Holotype 0 , UN- ITED STATES: California, Los Angeles Co., Los Angeles (USNM, type no. 1967). Condition of type: antennae incomplete; left middle leg missing except for coxa and trochanters; right hind tarsi and left hind tarsi 2-5 missing; right forewing detached and stuck to locality label;

SPECIMENS EXAMINED AND DIS-

ovipositor apex broken. 48 females and 42 males were examined (AEIC, CASC, CNCI, DBWC, UCDC, UCRC, USNM) from: UNITED STATES: California (Arroyo Seco Camp- ground in Monterey Co. , Ash Mountain in Sequoia National Park; Berkeley, Cabazon, Camp Baldy in San Bernadino Co. , Crystal Lake in Los Angeles Co., Devil’s Basin in El Dorado Co. at 8200 ft, Edgewood Park in San Mateo Co., Hetch Hechy Reservoir in Tuloumne Co., Los Angeles, Mill Valley, Mix Canyon summit in Solano Co., Petaluma, Pulga, Riverside, Rumsey, Santa Cruz Mts., San Francisco, Santa Ynez Mts., Snow Creek in Riverside Co., Tamarack Lake in El Dorado Co. at 7700 ft, Tanbark Flat in Los Angeles Co., Whitewater); Nevada (Haines Canyon in Douglas Co. at 5000 ft, Reno); Oregon (Chandler State Park in Lake Co. , Klamath Lake). Fig. 9 shows the distribution of this species. Collection records indicate that the species is found up to about 9000 ft. Dates of collection are usually between late May to July. The earliest date is 8 April 1978 (Yolo Co., California), the latest is 3-6 August 1965 (Marin Co., California).

COMMENT, Females have sometimes have large mesopleural punctures, separated by 0.5- 1 . 0 ~ their diameter. The amount of brownish red on the scape, pedicel and mandible is variable, ranging from almost completely brownish red, to fuscous with only hints of brownish red. Coxal varies, with brownish red sometimes on almost all the fore and middle coxae and the apical 0.3 of the hind coxa.

Cushman (1933) stated that male wing colour infuscipennis is clear, in contrast to the brown of, the female. This is not so, as examination of material identified by Cushman reveals that he mistook a male of whitei forfuscipennk.

The mature larva of fuscipennis is illfistrated in Fig. 5 . Cushman’s misidentification resulted in Short’s publishing a figure of whitei as fuscipennis (Short, 1959, 1978). Finlayson’s illus- tration of fusczpennis (1975) is actually that of pellogonia (see comment for that species).

A male from Edgewood Park (iv.1982, R . R. White; CASC) was reared from E.editha.

Benjaminia paeminosa sp.n. (Figs 18,27)

DIAGNOSIS. This species can be separated from others in thefuscipennis group by: setae of

15

Q9 18

21


16 17

22 23

FIGS 15-18. Dorsal aspects of heads: 15, B.euphydradis; 16, B.fuscipennis; 17, B.phae0thri.x; 18, B.paerninosa.

FIGS 19-26. Ventral aspect of sternite 8: 19, B.euphydryadis; 20, B.fuscrpennis; 21-23, B.pellogonia; 24- 26, B . whitei.

head and mesosoma white; hypostomal-man- 3. Supraclypeal area, and central region of dibular index=0.5; rugosopunctate meso- clypeus, rugosopunctate. 4. Gena in dorsal view pleurum; wings tinted with brown; tegula, and strongly receding, weakly convex (Fig. 18). 5 . legs beyond coxae, brownish red; punctures of Epicnemial carina not produced ventrally. 6. hind femur separated by 2-3X their diameter. Punctures of mesopleurum moderate to large,

MALE. Structure. 1. Hypostomal-mandibu- subadjacent, surface rugosopunctate. 7. Punc- lar index=OS. 2. Antenna with 34 flagellomeres. tures of anterior face of hind femur separated by

FIG. 17. Known distribution of Benjaminia spp,, B.carlsoni (half-shaded square). B.paeminosa (solid square). B.pellogonin (circles). B.phaeorhrix (triangle). FIG. 2 s . Known distrihution of B. ti.hirei.

3-3 X their diameter. 8. Surface of propodeuni coarsely rugosopunctate: carinae present except for sections of median longitudinal carinae apicad the basal transverse carina.

Coloitr.. Head and mesosoma black. the fo l - lowing brownish red: apex of mandibles. palpi, tcguln. extreme apices of coxae and remainder o f legs cxcept for brown hind tarsus. Wings with brown tint. Basal 0.7 of petiole fuscous; remainder of metasonia brownish red except for brown gonoforceps.

Lerigrh. 8.7 mm (approximately); forewing 6.3 mm.

TYPE MATERIAL. Holotype 6. UNITED STATES: California, Butte Co. . 15 miles E-NE chico. Mud Creek. ex E.c(fi iha. 1973 (White.) (USNM). Condition o f type: intact. except genitalia removed and in vial pinned with spccirnen.

ETYMOLOGY. From the Latin/~rrc~/iiiriosri.\. rough or uneven, in reference to the mesopleural sculpturing.

Benjaminia pellogonia sp.n. (Figs 6 , 21, 22, 23,27)

[ B r i i j ~ ~ i i i i i i i ( ~ jirsciperitiis Finlayson, 1975: 77. Misdetermination of larva]

DIAGNOSIS. Distinguished from other species in thefiiscipeririis group by: setae of head and mesosoma white; hypostomal-mandibular index= 1 .O : punctures of mesopleurum moderate. separated by 0.3-0.8x their diameter: wings tinted with brown: legs beyond second trochanter brownish red: punctures of hind femur separated by 2-3X their diameter; apicolateral corners of T7 fuscous.

FEMALE. Sfritcfitrc. 1. Hypostomal-mandibular index= 1.0. 2. Antenna with 32-35 flagellomeres. 3. Supraclypeal area centrally rugulosopunctate; punctures o f clypeus separated by 0.5-1 .Ox their diameter. on smooth surface. 1. Gena in dorsal view moderately receding and weakly convex, similar to Fig. 17. 5 . Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum moderate. separated


by 0 . 3 - 1 . 0 ~ their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 2-3X their diameter. 8. Surface of propodeum coarsely rugosopunctate; basal transverse and median longitudinal carinae complete, remaining carinae absent or indistinct.

Colour. Head and mesosoma black. the following brownish red: extreme apex of mandible. palpi. femora, tibiae, fore and middle tarsi. Sec- ond trochanters with varying amounts of brownish red, ranging from slight brownish tinge to almost completely dark brown. Hind tarsus brown. Tegula fuscous. Wings with brown tint. First metasomal segment with fuscous coloration varying from basal 0.7 of petiole, to all of petiole and basal 0.5 of postpetiole; remainder of metasoma brownish red except for transverse black marks on apicolateral corners of T2.

Lengrh. 8.4-9.3 mm (8.5 mm); forewing 5.7- 6.5 mm(6.9mm).

MALE. Structure. Similar to female; S8 as in Figs 21-23. Colour. As in female, except petiole always completely fuscous, fuscous of postpetiole varying from basal 0.3-0.8; T2 with median basal 0.2 fuscous; gonoforceps dark brown. Length. 7.5-8.5 mm; forewing 5.5-6.6 mm.

TYPE MATERIAL. Holotype 9 , UNITED STATES: Wyoming, Park Co., Beartooth Mountains, ex E.gillettii, vii. 1981 (Williams) (AEIC). Condition of type: intact. Paratypes, CANADA: Alberta, Lake Louise, 'No. A1530B; emerged 5 July 1950; F.I.S. 1930; lot 51-32', ex Arctiidae (19, CNCI); UNITED STATES: California, Alpine Co. , Ebbett's Pass at 8800 ft, ex Eeditha, 1972 (White) 119. USNM); Marin Co., Mill Valley, Blithedale Ridge, 24.iv.1972 (Amaud) (1 0, CASC); Napa Co., Butts Creek, ex E.editha luestherae, 15.iii.1974 (Reinhard) (1 8, AEIC); Santa Clara Co. , Alum Rock Park, with associated cocoons of unidentified nymphalid host, 14.i.1984 (Reinhard) (1 9, 16, CASC); Santa Clara Co., Palo Alto, 4.iii.1983 ( 1 8 , USNM); Idaho, Kootenai Co., 2200 ft ex E.colon, vi.1978 (White) ( 1 9 , AEIC); Latah Co., Moscow, 5.vi.1930 (Aldrich) (1 0 , USNM); Oregon, Klamath Co. , Klamath Falls, 7-24.v (Fox) (18, CASC); Wyoming, Park Co. , Beartooth Moun- tains, ex E.gilletti, vii.1982 (Williams) (1 d , NSIC); same data as preceding but collected vii.1981 ( 3 8 8 , AEIC); Sublette Co. , 7000 ft, ex Egillettii (White) (2 0 , 1 8 , AEIC; 2 9 , 1 8 , USNM).

COMMENT. This species is very similar to wliirei except for: ( I ) brownish red area of mandible confined to extreme apex: (2) consistently fuscous tegula; (3) more extensive brownish coloration of the second trochanter; (1) postpetiole usually with at least basal 0.3 fuscous: ( 5 ) fuscous apicolateral corners of T2 in pellogonin is the best character to separate the two species. The shape of S8 in pellogorzin is apically retangular (Figs 21-22) except for a male from Wyoming (Bear Tooth Mts.. AEIC; Fig. 24. setae not figured); whirei has the apex rather evenly convex (Figs 24-26).

Finalyson (1975: 77) illustrated a larval head capsule of this species (Alberta: Lake Louise, CNCI) as B.fusciperinis. She stated that it was 'reared from an unknown woolly Lepidopterous larva', although the label data with the adult reads 'ex Arctiid'. In view of the lack of rearing records from other hosts. the identification was probably incorrect.

ETYMOLOGY. From the Greek pellos. dusky. and goriia. angle, in reference to the fuscous coloration of T2.

Benjaminia pbaeotbrixsp.n, (Figs 17.27)

DIAGNOSIS. This species can be separated from others in the fusripennis group by: setae of head and mesosoma dark brown; hypostomal- mandibular index= 1 .O; rugosopunctate mesopleurum; wings brown; femora and tibiae dark brown; punctures of hind femur separated by about 3xtheir diameter.

MALE. Srrrrcture. 1. Hypostomal-mandibular index=0.4. 2. Antenna with 36 flagellomeres. 3. Supraclypeal area, and basal 0.5 of clypeus, rugosopunctate; apical 0.5 of clypeus with punctures separated by 0 . 3 - 0 . 5 ~ their diameter, on smooth surface. 4. Gena in dorsal view moderately receding, weakly convex (Fig. 17). 5. Epicnemial carina not ventrally produced. 6. Punctures of mesopleurum moderate to large, subadjacent to separated by about 0.2 X their diameter. surface rugosopunctate. 7. Punctures of anterior face of hind femur separated by about 3x their diameter. 8. Surface of propodeum strongly rugosopunctate. some rugae almost as produced as carinae; all carinae present and well developed.

Colour. Head and mesosoma black; apex of

296 David Wahl

smooth surface. 4. Gena in dorsal view moderately receding and weakly convex, similar to Fig. 17. 5. Epicnemial carina not produced ventrally. 6. Punctures of mesopleurum moderate and separated by about 0.3-0.8X their diameter, on smooth surface. 7. Punctures of anterior face of hind femur separated by 2-3x their diameter. 8. Surface of propodeum coarsely rugosopunctate: basal transverse carina and median longtidinal carinae usually complete, median longtiduinal carinae sometimes absent near apex, lateral longitudinal carinae rarely complete, remaining carinae absent or indistinct.

Colour. Head and mesosoma black, the following brownish red: apex of mandible, palpi, and legs beyond first trochanter. Tegula varying from completely brownish red to dark brown. Wing with brown tint. Basal 0.7-0.8 of petiole fuscous; remainder of metasoma brownish red.

Length. 9.7-11.8mm(9.9mm);forewing6.1- 7.8 mm (6.9mm).

MALE. S/rucfure. Similar to female; S8 as in Figs 24-26. Colour. As in female except petiole occasionally completely fuscous; gonoforceps dark brown. Length. 8.8-10.3 mrn; forewing 6.0-7.3 mm.

TYPE MATERIAL. Holotype 9 , UNITED STATES: Oregon, Josephine Co. , Selma, 26.v. 1978 (Townes) (AEIC). Condition of type: intact. Paratypes, UNITED STATES: Califor- nia, Alameda Co. , 11 miles SE Livermore on Mines Road, ex Euphydryas chalcedona, 1971. (White) (1 sex?, CASC); (Mariposa Co., Briceberg, ex E.chalcedona, 1971 (White) ( 3 9 , 1 sex?, CASC); Mono Co., Tioga Pass at 10,500 ft, ex E.editha nubigena, 26.vi-12.vii.1970 (Mol- denke) (18, USNM); Napa Co., Angwin at 1600 ft, 22.vi.1967 (Bauer) (1 sex?, CASC); Placer Co., ‘thro C.V. Riley 1888’, ( 1 6 , USNM); San Mateo Co., Jasper Ridge 1Stan- ford Biol. Preserve), ex E.chafcedona, 1973 (White) ( 1 9, AEIC); Santa Clara Co., 6 miles E- SE San Jose, Silver Creek, ex E.editha, 1971 (White) ( 3 9 , CASC); same data as preceding, but collected 1978 (16, USNM); Santa Clara Co. , 6 miles E . Morgan Hill, Coyote Reservoir, ex Eed i tha , 1971 (White) (16, USNM); same data as preceding, but collected 1972 (19, USNM); Stanislaus Co., circa 12 miles W. Pat- terson, Del Puerto Canyon, ex Eedi tha, 1971 (White) ( 1 9 , 16, AEIC); Idaho, Butte Co. , Craters of the Moon National Monument, 1 .viii. 1964 (Westcott) (9, USNM); same data as

FIG. 29. Metasoma of 6 .~3h i t e i

mandible and legs beyond first trochanters, dark brown. Wings brown. Setae of head and mewsoma dark brown. First metasomal segment with petiole and basal 0.5 of postpetiole fuscous; remainder of metasoma deep brownish rcd except for dark brown of apical 0 .7 of gonoforceps.

Lcngth. 10.3 mrn (approximately); forewing 6 . Y mm.

TYPE MATERIAL. Holotype 8, UNITED STATES: Idaho, Butte Co. , Craters of the Moon National Monument, Little Cottonwood Creek, ‘collecting station no. l ’ , on Heracleum lanatum. 8.vii.1965 (Homing) (USNM). Type condition: right hind leg beyond coxa missing; genitalia removed and in vial pinned with specimen.

ETYMOLOGY. From the Greek phaios. dusky or brown, and thrix. hair, in reference to the dark brown setae of the head and mesosoma. It is a n o u n in apposition.

Benjaminia whitei5p.n. (Figs 7,24,25.26.28)

[Benpmrnrrz fuscipennis Short, 1959: 588. Mis-

[Renjurninia firscipennis Short. 1978: 80. 352. determination of larva]

Misdetermination of larva]

DIAGNOSIS. Distinguished from other species in thefuscipennis group by: setae of head and mesosoma white; hypstomal-mandibular index= 1 .O: punctures of mesopleurum moderate. separated by 0.3-0.8x their diameter; wings tinted with brown: legs brownish red beyond first trochanter: punctures of hind femur separated by 2-3x their diameter.

FEMALE. SrrLic/ure: 1. Hypostomal-mandibular index= 1.0. 2. Antenna with 33-36 flagellomeres. 3. Supraclypeal area centrally rugulosopunctate; punctures of clypeus separated by about 0.5-1.Ox their diameter. on

Revision of Benjnminia 297

note of thanks is due R. R. White (Stanford Uni- versity) for sending his field notes associated with his collection of Berrjnrnirlin and for other assistance. Dimitri Kasparyan very kindly pro- vided translations of Russian locality labels. Mark R . Shaw (Royal Scottish Museum, Edin- burgh) was of great help in both loaning specimens and providing information about them. Ernest Williams (Hamilton College) donated his specimens of B.pellogonia to the American Entomological Institute collection and his generosity is gratefully acknowledged.

This research was supported by an NSF Post- doctoral Fellowship in Environmental Biology.

preceding, but collected 1 .viii. 1964, on Herac- leum lanatum (Horning) (2P . USNM); Kootenai Co., 2200 ft , ex E . colon (White) (1 9 , AEIC); Nebraska, Sioux Co., Munroe Canyon, 'em. 9 June 1982; ex Charidryas (Euphydryas)' ( 2 9 , VKGC]; Oregon, Josephine Co., Selma, 21.v.1978 (Townes) (18, AEIC); Washington, Pierce Co., Mt Rainier, Sunrise, 6318 f t (Wil- cox) (2 '2, USNM; l d , OSUO).

OTHER SPECIMENS. UNITED STATES: California, San Luis Obispo Co., San Luis Obispo, Madonna Inn, 1971 (Singe) (18, CASC); Santa Cruz Co., Santa Cruz Mts (1 sex?, USNM); Colorado, Park Co., Fairplay, ex Eanicia, vii.1978 (White) (1 6, USNM).

COMMENT. The use of S8 to separate whitei males from those of pellogonin is discussed in the description of the latter species.

A few specimens are found with the hind second trochanter basally dark brown. The two females from Nebraska have a deeper brownish red coloration of the legs and metasoma than in other specimens of this species.

Two specimens are tentatively assigned to this species. One male from California (San Luis Obispo) has the mesopleural punctation more crowded and rugosopunctate in appearance, the maxillary palpi dark brown, and the propodeum with only the basal transverse carina present on a more delicately rugosopunctate surface than i s normal for whitei. S8 is typical for whifei. Poor preservation of the specimen precludes any further analysis. The specimen from Colorado (Fairplay) is morphologically identical to typical whifei but the metasoma exhibits more fuscous coloration on TS-8. as well as on S1-8 (Fig. 29). This colour pattern. as well as its isolated location relative to the rest of the species. leads me to place i t in t+,hitci with a query.

The specimen from the Santa Cruz Mountains of California is the source of the larval remains referred to as B.fi~scipennis by Short (1959).

ETYMOLOGY. This species is named for R. R. White. in recognition of collecting Berrjnniirrin specimens as part of his butterfly research.

Acknowledgments

1 thank Henry Townes and Virendra Gupta (American En tomological lnsti t u tc) for their advice and criticisms during thc study. A special

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Accepted 30 August 1988

Documents

A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)