D. Christopher Darling.
Description of a New Species of Krombeinius (Hymenoptera: Perilampidae) from the Philippines, and the Phylogenetic Relationships of the Genus.
Psyche 89(3-4):307-316, 1982.

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DESCRIPTION OF A NEW SPECIES OF
KROMBEINIUS (HY MENOPTERA: PERILAMPIDAE) FROM THE PHILIPPINES, AND THE PHYLOGENETlC RELATIONSHIPS OF THE GENUS*
BY D. CHRISTOPHER DARLING
Department of Entomology,
Cornell University, lthaca, N.Y. 14853
The genus Krombeinius (Hymenoptera: Perilampidae) was re- cently described (BouEek 1978) to include perilampids with an amalgam of the characters of Euperilampus Latreille and Peri/am- pus Walker. The habitus suggests Euperilampus, and there are two synapomorphies to unite these two genera (Darling 1983): postspi- racular sclerite reduced to a narrow triangle, less than one-half as wide as the adjacent pronotum; and pronotum massive, at least one-third the length of the mesoscutum. However, Krombeinius exhibits the wing venation, presence of a marginal rim on the scutel- lum, and large third metasomal tergite characteristic of Perilampus. I regard these as plesiomorphic similarities. The genus is character- ized by the absence of the defining apomorphic characteristics of Euperilampus, i.e., by symplesiomorphy.
In this paper I present new information on the structure of the male genitalia and labrum of the type species of Krombeinius. These structures have proved to be of considerable value in defining gen- era in the Perilampidae (Darling 1983). From this analysis I suggest autapomorphies for defining Krombeinius. In addition, 1 describe a new species of Krombeinius from the Philippines, and discuss the affinities of the three included species. Taxonomic studies of Krombeinius have been hampered by the scarcity of material. The type species, K. eumenidarum, was de- scribed by BouEek (1978) from a series of specimens (2 male, 2 female) reared from the larvae of an eumenine wasp in Sri Lanka. All specimens were prematurely killed and had to be liberated from the pupal cuticles, producing some abnormalities in the type mate- rial. Also included by Boutek (1978) in this genus was Perilampus megalaspis Cameron, known only from the type material (3 females) *Manuscript received b.v the editor September 22, 1982.



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308 Psyche [VOI. 89
and an additional female, all from Sarawak, Malaysia. During my study of material in the U.S. National Museum of Natural History, Washington [USNM], I located an additional male specimen of K. eumenidarum. This specimen [India: Kerala Survey, 12.5 Pechipa- rai, 25-27 August 19741 was dissected and re-mounted and is the basis for the description of the labrum and male genitalia. In addi- tion, the C. F. Baker Collection [USNM] contained a new species of Krombeinius from the Philippines, which I describe herein. Abbreviations used in text: Fl-7: funicular segments 1-7; MSC: length of mesoscutum along midline; OOL: length of ocular-ocellar line; PN: length of pronotum along midline; POL: length of poste- rior ocellar line; SC: length of scutellum along midline; T2-8: meta- soma1 tergites 2-8.
Krom beinius
Krun7heiniu.s Boucek. 1978: 302, Figs. 1.2. Type species: Krombeinius eumenidarum Boucek, 1978: 302, Fig. 1. [original designation].
Diagnosis:
Hymenoptera: Chalcidoidea: Perilampidae (sensu Graham, 1969). Species of Krombeinius can be distinguished from Monacon Water- ston, Burksilampus Bouiek, Stejfanolampus Peck and Perilampus Latreille by the narrow postspiracular sclerite, less than one-half the width of the adjacent pronotal panel, and from Euperilampus by having the marginal vein longer than the postmarginal (Fig. 1). All known species are moderately large, 3 to 5% mm long, black without metallic reflections and are restricted to the Oriental region. There are three species currently placed in Krombeinius: K. eume- nidarum Boucek, K. megalaspis (Cameron) and K. saunion, nsp. A revised key to the species of Krombeinius is not presented. The key of Boucek (1 978) separates K. eumenidarum and K. megalaspis. An additional character to separate these two species is the inner orbits: costate in K. eumenidarum (Fig. 8), and smooth in K. meg- alaspis (Fig. 12). K. saunion is readily distinguished from these two species by the prominent spine at the apex of the scutellum (Fig. 1,15). The apex of the scutellum is truncate in the other two species (Figs. 7,ll).




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19821
Darling- New Species of Krombeinius
Fig. 1, Krutnbeinius saunion, lateral habitus. Description:
Head: supraclypeal area smoothly convex, without horn or ridge; scrobal cavity deep, extending to lower ocular line or to middle of clypeus; clypeus and supraclypeal area separated by distinct suture or by faint line; inner orbits carinate; frontal carina separating the median and posterior ocelli; malar sulcus absent; malar region with strong oblique costae; posterior ocellus located high on vertex, POL approximately equal to OOL; labrum with a single narrow stalk, expanded distally with 7 digits, each with a terminal seta, and with pair of sessile setae not associated with digits, strongly excised medially [n = 1, K, eumenidarum, Fig. 31. Mesosoma: dorsum of pronotum smoothly convex, without transverse elevations; pronotum massive, about one-third length of mesoscutum, not narrowed medially; mesothoracic spiracle located between pronotum and sidelobe of mesoscutum; postspiracular sclerite fused to the pronotum but delimited by surface sculpture; postspiracular sclerite less than one-half width of adjacent pronotal panel, with many or a single puncture; notauli distinct; scutellum vaulted, jutting over propodeum and base of metasoma; apex of scutellum acuminate, or truncate or with a distinct spine; propo- deum with median area foveate, or with a short median ridge, sub-



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Psyche
Figs. 2-5. 2.3 Krumheinius eumenihrum. 2. Male genitalia. Inset: enlargement of paramere. 3. Labrum. 4.5. Euperilampus triangularis (Say). 4. Apex of male genita- lia. 5. Labrum. [Scale lines 0. I mm.]




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19821 Darling- New Species of Krombeinius 311 median areas with weak transverse rugae or aciculate; basitarsomere not conspicuously lengthened. Forewing with marginal vein longer than postmarginal, postmarginal vein long, about 3 times length of stigmal vein, stigmal vein making either a right or oblique angle with marginal vein.
Metasoma: petiole short, transverse, the tergum forming a ridge along anterior face of gaster, sternum shifted posteriorly; gaster triquetrous, T2 and T3 fused, covering most of dorsum; T2 without distinct basal fovea; T3 much longer than T2, subquadrate, slightly wider than length along midline; ovipositor ventral, not upturned, sheaths not distinctly exserted; male genitalia with distinct para- meres [n = 1, K. eumenidarum, Fig. 21.
Discussion:
The male genitalia of Krombeinius eumenidarum (Fig. 2) are similar to those of species of Perilampus: the parameres are distinct, and strong setae are distributed on these lobes. This configuration occurs throughout the Chalcidoidea (see Domenichini 1953) and is regarded as plesiomorphic. In Euperilampus a derived condition is found (Darling 1983): distinct parameres are lacking, and the basi- paramere has a patch of strong setae distributed on transparent areas laterad of the ventral lobe (Fig. 4). The labrum of Krombeinius eumenidarum (Fig. 3) has a narrow central stalk, not found in other perilampid genera (Riek 1966; Domenichini 1969; Darling, unpublished). However, the labrum does share synapomorphies with species of Euperilampus (Fig. 5) including a reduced number of digits (7 or 8), a pair of smaller, sessile setae not associated with digits, and a strong median exci- sion. The narrow stalk distinguishes the labrum of Krombeinius from those of Euperilampus, and is postulated as an autapomorphy of Krombeinius. All other perilampid labra are 10-12-digitate, and not as strongly excised medially.
The host association of the type species of Krombeinius [larva of Vespidae: Eumeninae] is different from that of any other described perilampid, although solitary Sphecidae are attacked by some Peri- lampus species (e.g., Perilampus nitidus, primary parasitoid of Ectemnius paucimacwlatus. Krom bein 1964, as P. canadensis). This behavioral character is regarded as an additional autapomorphy for the genus Krombeinius.




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Psyche [VO~. 8
Figs. 6 13. 6 9. K1~01iiheit7iu.s ~unn'nu~~uni. 6. Head, dorsal. 7. Mesosoma, dor- sal. 8. Head, frontal. 9. Head, lateral. 10- 13. A.'. mqplu.spni. 10. Head, dorsal. I I. Mesosoma, dorsal. 12. Head, frontal. 13. Head, lateral. [Scale line 1 mm.]



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Darling - New Species of Krombeinius
Krombeinius sauion n.sp.
(Figs. 1, 14- 17)
Type Locality: Philippines, Mindanoa, Surigao. Type Material: Holotype: Female [Baker Collection, USNM]. Etymology: The specific epithet is a noun in apposition, Greek for "javelin", and is a reference to the elongate spine on the scutellum of this species.
Diagnosis:
This species can be immediately recognized by the prominent spine at the apex of the scutellum (Figs. 1, 15). The apex of the scutellum is truncate in K. eumenidarum (Fig. 7) and K. megalaspis (Fig. 1 1).
Description:
Female: Length, 5.4 mm. Black, except tegula and flagellum brown, mandible reddish-brown, apex of foretibia and spur, and tarsi yellow; wings hyaline, veins darkened. Head: length of malar space 0.34 eye height; OOL 0.95 POL; maximum width of scrobe 0.56 head width; head transverse, width: height = 1.17; inner and outer orbits costate, costae convergent on clypeus; scrobal cavity deep and wide, extending below lower ocular line and delimiting clypeus and supraclypeal area; clypeus trans- verse, width:height = 1.31, polished and covered with long setae; clypeus not delimited by sutures, upper margin indicated by a faint line, tentorial pits distinct, lower margin weakly emarginate; ocular- ocellar region with costae radiating from posterior ocellus; vertex with strong costae at posterior margin; supraclypeal area glabrous, height 0.5 1 clypeus height; lateral wall of scrobe merging smoothly with face; lower tooth of mandible pointed at apex; base of mandi- ble with weak punctures; labio-maxillary complex short. Antenna: scape narrowly linear, length 8.5 maximum width; pedicel and funicular segments subequal in length (1 8 versus 14,15,17,16,15, 15,13;F1-F7); pedicel 0.21 scape length; anellus 0.43 length of F 1 ; Fl elongate, remaining flagellomeres transverse; clava 0.25 length of funicle.
Mesosoma: pronotum massive, PN:MSC = 0.34, lateral pronotal collar not regularly convex, suggesting bumpy shoulders; scutellum acuminate with a long spine, SC:MSC = 1.75; dorsum of pronotum



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314 Psyche [vo~. 89
punctate-reticulate, punctures coalesced to form weak irregular transverse costulae medially; midlobe of mesoscutum and scutellum weakly punctate, becoming punctate-reticulate along notauli; side- lobe of mesoscutum smooth along notauli, laterally punctate- reticulate; notauli distinct; scutellum in lateral view tapering abrupt- ly towards apex (Fig. 1 ; cf. Figs. 1,2 in BouEek 1978); underside of scutellum mostly coriarious, with shallow convergent grooves; propodeum vertical, medially about twice as long as metanotum, with weak median ridge, submedian areas with transverse costulae, callus reticulate-rugose; width of postspiracular sclerite 0.44 width of adjacent pronotal collar, with about 10 foveae; axilla reticulate above, costate below; axillula smooth. Forewing: submargina1:mar- gina1:postmarginal:stigmal veins as 64: 17: 12:4; stigmalvein making a right angle (90 degrees) with marginal vein. Metasoma: T2 smoothly concave with weak coriarious sculpture; T2 with sparse setae, without punctures, border between T2 and T3 indicated by a suture, laterotergite glabrous; T3 massive and con- vex, about twice length of T2 along midline, length about equal to maximum width (22 versus 25), evenly covered with long setae except along T2 border and along margins of tergite, without punctures.
Male: Unknown.
Discussion:
Krombeinius saunion is more closely related to K. eurnenidarum than to K. megalaspis. Synapomorphies of these two species are: the stigma1 vein making a right angle with the marginal vein (oblique in K. megalaspis and outgroup: Euperilampus and Perilampus); cly- peal-supraclypeal margin weak or indistinct (separated by distinct suture in K. megalaspis and outgroup: Euperilampus and Perilam- pus); lateral pronotal collar suggesting bumpy shoulders (regularly convex in K. megalaspis and outgroup: Euperilampus and Perilam- pus); and postspiracular sclerite with many foveae (a single fovea is found in K. megalaspis, and in the ancestral species groups of Eupe- rilampus, Darling 1983). Considering Euperilampus as the out- group, the following similarities of K. eumenidarum and K. saunion are regarded as plesiomorphic: propodeum medially about twice as long as metanotum (equal to metanotum in K. megalaspis; autapo- morphy); scutellum, in lateral view, not strongly convex, tapering gradually towards the apex (highly convex in K. megalaspis, Boucek 1978: Fig. 2; autapomorphy).




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19821 Darling- New Species of Krornbeinius 3 15 K. saunion and K. eumeniciaruni also have the inner orbits with strong costae (Figs. 8,9,16,17), whereas the inner orbits of K. meg- alaspis are smooth (Figs. 12,13). 1 consider the costate inner orbits to be a synapomorphy of Eupe~~ilanipus + Kromheinius. As such I interpret the smooth orbits of K. megalaspis as an autapomorphic reversal. A similar reversal in this character is indicated in the Eupe- rilai?ipus cladogram (Darling 1983).
There remain some difficulties in justifying the current composi- tion of the genus Kromheinius. The numerous characters separating Figs. 14- 17. Krontheinius suut?ion. 14. Head, dorsal. 15. Mesosoma, dorsal. 16. Head, frontal. 17. Head, lateral. [Scale line 1 mm.]



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316 Psyche [vo~. 89
A'. eumenidaruin + K. suunion from A'. megu/aspis question the inclusion of A'. megalaspis. A revised classification would create a monobasic new genus for Peri/u/~ipus mega/aspi.s, and would allow A'rombei~ius to be defined by the synapomorphies of K. eumenida- rum + K. saunion. Recalling that the proposed synapomorphies of Kron1beiniu.s (structure of the labrum; host association) are not known for K. niega/uspis, it would not be surprising if this species were to be excluded at some later date. Clearly, more material and associated biological information are essential to re-evaluate the composition of Kro~nheinius, and any nomenclatural changes at this time would be premature.
1 would like to thank the following people for comments on the manuscript: W. L. Brown, Jr., J. M. Carpenter, G. C. Eickwort, S. W. Nichols, and Q. D. Wheeler.
The habitus drawing was skillfully prepared by Jim Miller. This research was supported in part by a National Science Foun- dation Dissertation Improvement Grant.
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DARLING, D. C.
1983. A review of the New World species of Euper//ut~/)us (Hymenoptera: f'erilampidae), with notes about host associations and phyloge- netic relationship;). In pre.\.\. Quaestiones entomologicae. DOMI..SKHIM, G. 1953. Studio sulla morphologia dell'addome delgi Hymenop- tera Chalcidoidea. Bollettino di Zoologia Agraria e Bachicoltura, 19: 183-298, 27 t'ii",.
D~MI.\KHIM. G. 1969. Material! per la morfologia comparata degli Hymenop- tera Chalcidoidca. Memorie dclla Societa Entomologica Italians, 48:584-608, 53 ligs.
GRMIAM. M. W. R. DI: V. 1969. The Ptcromalidae of north-western Europe (Hymenoptera: Chalcidoidea). Bulletin of the British Museum (Natural History) Entomology Supplement 16. 908 pp. KR~MIWIN, K. V. 1964. Natural History of Plummers Island, Maryland XV111. h e hibiscus wasp. an abundant rarity, and its associates (Hymenoptera: Sphe- cidae). Proceedings of the Biological Society of Washington, 77:73 I 12.
RII.K. E. F. 1966. Australian Hymcnoptera Chalcidoidea, Family Ptcromalidae, Subfamily Pcrilampinae. The Australian Journal of Zoology, 14: 1207 - 1236.



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