PROC. ENTOMOL. SOC. WASH. 107(3). 2005, pp. 576-595 REVISION OF THE NORTHERN SOUTH AMERICAN TIPHIID GENUS MERITHYNNUS KIMSEY, 1991 (HYMENOPTERA: TIPHIIDAE: THYNNINAE) Lynn S. Kimsey Bohart Museum of Entomology, University of California, One Shields Ave., Davis 95616, (e-mail: [email protected]) — Abstract. The South American genus Merithynnus is revised. Five new species from Colombia, M. bilobatus, M. intermedius, M. montanus, M. nasalis and M. paradoxus, are described. Of these species only paradoxus and intermedius are known from both sexes. The remaining species are based on males. Seven previously described species, three from Venezuela (osorioi, pecki, and turbulentus) and four from Colombia (acumi- natus, bogotanus, desertus, and mimulus) are redescribed, and a distribution map and key to the species are provided. Merithynnus desertus is known only from the female and acuminatus, bogotanus, mimulus, and turbulentus are known only from males. Key Words: Merithynnus, Thynninae, Elaphropterini, Colombia For a number of years Michael Sharkey, The remainder of the previously described University of Kentucky, in collaboration species, osorioi, pecki, and turbulentus, are with the Alexander von Humboldt Biolog- from Venezuela. In the new Colombian ma- ical Resources Research Institute, Bogota, terial the type series ofparadoxus alone is has intensively collected Hymenoptera in represented by more than 200 specimens. parks across Colombia. This project has re- Although Merithynnus species are undoubt- vealed a heretofore unknown fauna ofwasp edly parasitoids, like other members of the species, particularly in the tiphiid subfamily family, nothing is known ofthe natural his- Thynninae. Only two genera of Thynninae tory of the genus. are recorded from Colombia, Merithynnus and Aelurus (Rhagigasterini). All of the Materials and Methods Colombian thynnines in the tribe Elaphrop- Specimens used in this study were bor- terini belong to the genus Merithynnus. rowed from the following museums and in- This study increases the number of known stitutions. Type depositions are indicated in Merithynnus species from seven to twelve. the text by the city name of the re—spective Oddly, none of the previously described museum in capital letters: BERLIN Hum- species from Colombia were represented in boldt Museum, Humboldt-Universitat, Ber- the material collected by Sharkey and the lin, Germany (F Koch); BOGOTA—Alex- Humboldt Institute. ander von Humboldt Biological Resources Prior to the study of Merithynnus in Co- Research Institute. Bogota—, Colombia (D. lombia, the genus was known from fewer Arias); BUENOS AIRES Division Ento- than 60 specimens, in most species just mologia, Museo Argentino de Ciencias Na- from the type series. Four of these, acumi- turales, Buenos Aires, Argentina (M. J. Vi- natus, bogotanus, desertus, and mimulus, ana); CAMBRIDGE—Museum ofCompar- were described from Bogota, Colombia. ative Zoology, Harvard University, Cam- VOLUME 107, NUMBER 3 577 USA bridge,—Massachusetts, (P. Perkins); ventral lobe or dorsal one, before apical DAVIS Bohart Museum of Entomology, strap, apica—l strap bending dorsally. University of California, Davis, USA (S. L. Female. Mandible slender, edentate; Heydon); GAINESVILLE—Florida State propleuron elongate and strongly convex Collection ofArthropods, Gainesville, USA ventrally; scutum and metanotum obscured; LONDON— (L. A. Stange); The Natural scutellum elevated and ovoid dorsally; me- History Museum,—London, England (S. tasomal tergum I strongly narrowed anteri- Lewis); OTTAWA Hymenoptera Section, orly with short, often densely setose, ante- Biosystematics Institute, Agriculture Cana- rior surface; tergum II with two strong da, Ottawa (L. Masner); OXFORD—Hope transverse ridges, apical margin strongly Entomological Collections, Oxford Univer- upcurved; pygidium coarsely punctate, with sity Museum, England (C. O'Toole). Pri- lateral carina, usually with one or more api- mary types were studied for all species. cal lobes orteeth, tufts ofsetae laterally and Modifications ofthe head, mesosomaand one or more on the surface of the plate and terminalia are most important to distinguish on apical margin; sternum VI apical plate species. In females the shape ofthe protho- broadly ovoid o—r forming half circle. rax, propodeum and pygidium are most sig- Distribution. Species of Merithynnus nificant. Diagnostic features of males in- appear to be restricted to higher elevations clude the configuration of different com- of the northern reaches ofthe Andes in Co- ponents of the face, body coloration, and lombia and Venezuela, in the Cordillera the shape ofthe hypopygium, paramere and Oriental and Cordillera Merida (Fig. 1). aedeagus. The terms terga and sterna are None of the specimens examinemd for this study were found below 1,000 and the used to refer to segments of the metasoma. majority of specimens were from between The slender, elongate, curved apex of the m m aedeagus is referred to below as the "apical 2,000 and 3,—000 in elevation. Discussion. The genus Merithynnus strap." Puncture diameter, given in the text was described by Kimsey (1991) to include as PD, is used to describe the density of a variety of species, some of which were punctation of the integument. originally described in Scotaena and Ela- Merithynnus Kimsey phroptera. Merithynnus is characterized by a number offeatures, some shared with oth- Merithynmis Kimsey 1991:74. Type spe- er members ofthe Tribe Elaphropterini. and cies: Merithynnuspecki Kimsey 1991:75. some unique to the genus. Unique features Original designation. include the configuration of the female py- — gidium, with lateral carina, irregularly Male. Mandible bidentate, usually with sculptured posterior surface, 1-4 apical small angle or tooth subbasally on dorsal lobes or teeth, and two or inore tufts of se- margin; pronotal transverse anterior ridge tae, male scutellum with septate, transverse laterally notched; scutellum with trans- anterior groove, male hypopygium apically versely septate groove along anterior mar- hoof-like, lateral margin with submedial gin; epipygium apically truncate to round- notch or indentation, and paramere apically ed, with lateral carina; hypopygium notched bending inward. Females are onl\ known submedially, ligulate or horse hoof-shaped for six species: hoi^otiuuis. dcscrtus, intcr- apically; paramere elongate, bending ven- inedius, osorioi, pecki, and parachixus. The trally along longitudinal axis, with apical female of desertus is uiidcnibtedly synony- tooth or lobe bending toward midline vol- mous with one o{ the other species de- sella appressed against aedeagus, with short scribed by Turner (1910) based on males broadly rounded to slender, free apical lobe; Worn Bogota, Colombia, but this cannot be aedeagal column short, often with large resolved without collecting pairs in copula. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON 578 Fig. 1. Distribution map ofMerithynnus species in Colonnbia and Venezuela. The genus is redescribed in light of phylo- 4. Pronotal medial disk strongly excavate, al- genetic analyses ofthe genera ofThynninae most trough-like anteromedially, excavation planai"withanteriorcollar(Fig. 24);pygidium currently underway. with carina enclosed apical snout or plate Key to the Species of Merithynnus (Fig. 33) .... intermedius Kimsey, new species - Pronotum medial disk convex or flattened, 1. Wingless (female) 2 strongly to somewhat elevated above anterior - Winged (male) 6 collar; pygidium without carina enclosed api- 2. Pronotal disk anterioriy trilobate (Fig. 32); cal snout or plate 5 pygidium with two subapical lobes or teeth 5. Propodeum saddled in profile (Fig. 37); pron- between apical tufts of setae (Fig. 35); pro- otum with medial disk convex, strongly ele- podeum abruptly concave adjacent to metan- vated above anterior collar (Fig. 37); pygidi- otum (Fig. 32) um apically quadrilobate (Fig. 36) paradoxus Kimsey, new species pecki Kimsey - Porrloynot(aasl idniskFigbsi.lob2a4t-e,25c,on3v0,ex37o)r;flpaytgaindteiruim- - Propodeum convex in profile, not saddled emarginate or unilobate between apical tufts (Fig. 25); pronotum nearly flat, medial disk ofsetae (Figs. 29, 33-34); propodeumstrong- only slightly elevated above anterior collar ly convex adjacent to metanotum (Figs. 24, (Fig. 25); pygidium apically trilobite, with 25, 37) 3 acute medial lobe (Fig. 29) . . desertus (Turner) 3. Propodeum dorsomedially dentate (Fig. 30); 6. Clypeus with medial lobe, tooth orprojection metasomal sternum VI apically forming half (as in Figs. 3, 10); metasoma with extensive circle (Fig. 34) osorioi Kimsey orange or reddish markings 7 - Propodeumdorsomediallyrounded(Figs.24- - Clypeus without medial tooth or projection 25, 37); metasomal sternumVI apicallysemi- (as in Figs. 4, 11); metasoma without orange circular (as in Figs. 29, 33, 36) 4 or reddish markings, except mimulus 9 VOLUME NUMBER 107, 3 579 Fig. 2. Side view ofmale and female Merithynnusparadoxus. Forecoxa with elongate ventral spine (Fig. subapically across subsidiary tooth (as in Fig. 13); clypeus apicomedial margin forming 9); clypeal apical projection with apex broad- elongate, narrow, upcurved lobe(Fig. 10);an- er than (as in Figs. 9, 11) or less than half tennal lobe sharply angulate apically (Fig. 10) distance between antennal sockets (Fig. 4) 10 paradoxus Kimsey, new species 10. Clypeus apicomedial projection with sides Forecoxa without ventral spine; clypeal api- convex to lobate, apex narrowly bidentate comedial margin unilobate, trilobate, or (Fig. 4); forewing marginal cell more darkly broadly truncate (as in Figs. 3-9, 11-12); an- stained, noticeably darker than rest of wing tennal lobe rounded 8 bilobatus Kimsey, new species Clypeus apical margin with small medial lobe - Clypeus apicomedial projection with sides and small obtuse sublateral lobes (Fig. 3) . . straight to concave, apex broadly bilobate, acuminatus (Turner) flat, bidentate or indented (as in Figs. 7, 9, Clypeus apicomedially slightly produced and 1 1); forewing marginal cell at most slightly flat medially, with small acute to obtuse sub- darker than rest of wing membrane II medial angles (Fig. 8) 11. Clypeus convex medially, rounded apicome- nasalis Kimsey, new species dially; mandible broadest submedially; face Metasoma and legs extensively reddish; man- with continuous yellow stripe along innereye dible broadest submedially (Fig. 6); clypeal margin connecting behind ocelli, forming apical projection with apex equal to distance continuous stripe between antennal sockets (Fig. 6) intertncdius Kimsey. new species initnulus (Turner) Clypeus flat medially, apicomedially angu- Metasoma and legs without reddish colora- late, not rounded in front view; mandible tion, black and yellow; mandible broadest broadest suhbasallv (as in Figs. 5, 9, 1 1); face X 580 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON with discrete yellow stripe along inner eye lobe and obtuse submedial ones; area be- margin not extending beyond ocelli 12 tween antennal socket and inner eye margin 12. Clypeus with apicomedial angles obtuse to with tiny punctures, punctures 1-3 PD 90° (Figs. 7, 11); mandibleslender, withsmall subapical tooth, broadest subbasally, without apart; frons with punctures larger than low- subbasal angle on inner margin (Figs. 7, 11) er face, nearly contiguous, somewhat elon- 13 gate; vertex punctures smaller than frons, Clypeus with apicomedial angles acute (Figs. contiguous; hindocellus 3 hindocellar di- 4, 10, 13); mandible with large subapical ameters from eye margin; flagellomere I tooth, broadest submedially, with small sub- basal angle on inner margin (Figs. 4, 10, 12) 2.5X as long as broad; flagellomere II 3.5 14 as long as broad; flagellomere XI 6X as 13. Clypeus flat in lateral view (Fig. 22); mandi- long as broad; mandible broadest medially, ble without subbasal angle (Fig. 11); aedea- with long apical tooth, large subsidiary gus withlarge ventral lobebeforeapical strap tooth and strong medial angle. Mesosoma: (Fig. 46) pecki Kimsey - Clypeus saddle-shaped in lateral view (Fig. Legs unmodified; pronotal punctures tiny, 18); mandible with obtuse subbasal angle 2-3 PD apart; scutal punctures 2-4 PD (Fig. 7); aedeagus without ventrallobebefore apart medially, becoming denser laterally, apical strap (Fig. 52) 0.5-1.0 PD apart; scutellar punctures 1-3 montanus Kimsey, new species PD apart; metanotum polished, punctures 14. Cgilnyaptee,uswiatphicaocmuetdeilaaltemraalrgainnglessha(lFliogw.ly9);emparro-- 2-5 PD apart; propodeum densely, trans- podeum finely shagreened; frons withoutyel- versely shagreened, punctures obscure, 2-4 low spot between ocelli osorioi Kimsey PD apart; mesopleuron densely, finely sha- - Clypeus apicomedial margin deeply emargin- greened, punctures 1-2 PD apart. Metaso- ate, with narrow, slightly up-turned lobes on ma: Finely shagreened, punctures tiny, 3- peiotdheerusmidhei(gFhilgy. p1o2l)isohredr;oufnrdoends(wFiigt.h5)y;elplroow- 5 PD apart; epipygium lateral carina strong- spot between ocelli 15 ly produced; hypopygium ligulate, apically 15. Clypeus apicomedial projection with lateral rounded, roughened dorsally (Fig. 26). margin obtuse or rounded (Fig. 5); face with Genital capsule (Figs. 38, 48): Paramere complete yellow stripe along inner eye mar- subtriangular, ventrally curved, tapering gin, extending to vertex behind ocelli; terga I-V with bright yellow stripe or spot; fore- apically in lateral view, ventral surface wing marginal cell only slightly darker than strongly concave, broad, apically subtrun- rest ofwing bogotanus Kimsey cate, inner angle slightly hooked in dorsal - Clypeus apicomedial projection with lateral view; aedeagus with large dorsal lobe. Col- margin straight (Fig. 12); face with two nar- or: Black, with yellow to red markings: rowly to well-separatedyellowish spotsalong eye margin, dorsal spot barely extending to clypeus ventrally yellow, antennal lobe yel- ocelli, not beyond; terga I-IV with small low, mandibles mostly yellow, inner eye whitish lateral spot; forewing marginal cell margin with yellow stripe extending to ver- markedly darker than rest of wing tex, connecting to stripe extending from turbulentus (Turner) posterior eye margin; pronotum with broad, Merithynnus acuminatus (Turner) transverse medial yellow stripe, large yel- (Figs. 3, 14, 26, 38, 48) low lateral spot; scutum with large, trilobate medial yellow spot, two small lateral spots Elaphroptera acuminata Turner 1910:194. and small yellow spot anteriorly on notau- Syntype males: Colombia: Bogota (BER- lus; scutellum with large yellow medial and LIN). — smaller lateral spot; metanotum yellow me- Male. Body length 14-15 mm. Head dially and laterally; propodeum with broad (Figs. 3, 14): Clypeus impunctate ventrally, U-shaped sublateral yellow spots, narrowly dorsally punctures contiguous to 0.5 PD separated medially, interrupted laterally in apart, with small medial tubercle, apical some specimens, with yellow dorsolateral margin barely produced, with acute medial spot; mesopleuron with large, J-shaped, yel- VOLUME 107, NUMBER 3 581 12.liirhulenlus Figs. 3-13. 3-12, Front view of male face, antennae removed. 13. Male foreleg. 582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON low dorsal spot and smaller ventral one; fo- large submedial tooth and subbasal obtuse recoxa anteriorly yellow; mid and hindcox- angle; flagellomere I 2.5X as long as broad; ae yellow dorsally and laterally; fore- and flagellomere II 3.5X as long as broad; fla- midfemora yellow ventrally; tibiae, tarsi gellomere XI 6X as long as broad. Meso- and hindfemur red to orange; metasomal soma: Pronotum weakly punctate, punc- tergum I yellow, with dark brown medial tures minute, 3-8 PD apart; scutum pol- spot and subapical orange spots; segments ished, punctures 1-4 PD apart; scutellar II-IV primarily yellow, with orange basal punctures 1-4 PD apart, laterally, nearly markings and orange to dark brown apical impunctate medially; metanotum impunc- markings; segment V blackish, with large, tate; propodeum finely shagreened, nearly lateral, yellow spot; segment VI black, with impunctate posteriorly; mesopleuron with lateral yellow tergal spot; epipygium dark dorsal margin of scrobal sulcus bulging, brown, with basolateral yellow spot; wing punctures 1-2 PD apart; grooves above pet- membrane yellow-tinted, veins yellowish iolar socket faintly septate; legs unmodi- brown. Ves—titure: Pale golden. fied. Metasoma: Terga and sterna finely Female. Unknown. shagreened, punctures shallow, 1-4 PD — Material examined. COLOMBIA: Bo- apart; hypopygium broadly rounded apical- gota (the type—series). ly, dorsal surface transversely wrinkled Discussion. Based on the coloration of (Fig. 27). Genital capsule (Figs. 39, 49): the male and modifications of the male Paramere slender, elongate, tapering apical- clypeus acuminatus is most similar to the ly in lateral view, broadened and slightly new species nasalis and paradoxus, de- hooked in dorsal view, with submedially scribed below. All ofthese species have ex- bulging inner margin; aedeagus with large tensive orange coloration on the legs and dorsal lobe. Color: Black, with pale yel- metasoma, and the clypeus has a medial lowish markings on lower two-thirds of tooth or projection. Merithynnus acumina- clypeus, antennal lobe, base of mandible, tus lacks the spined forecoxa and strongly face along eye margin with large spot ven- elongate clypeus of paradoxus. It can be trad and small spot dorsad, vertex with distinguished from nasalis by the clypeus transverse stripe, often interrupted subme- having a single apicomedial lobe and ob- dially, gena with large spot along lower eye tuse sublateral ones. margin, mandible pale yellow medially; pronotum with transverse submedial stripe Merithynnus bilobatus Kimsey, and large lateral spot; scutum with small new species medial spot and lateral stripe; scutellum (Figs. 4, 15, 27, 39, 49) with two medial and one anterolateral — Male. Body length 10-14 mm. Head spots; metanotum yellow medially, with an- (Figs. 4, 15): Clypeus flat in side view, ap- terolateral stripe; mesopleuron with com- icomedially elongate, with bilobate apex, ma-shaped dorsal spot, medial spot and side of elongation strongly convex, punc- posterior one; propodeum with lateral spot tures sparse and 4-6 PD apart apically, be- and submedial longitudinal stripes; coxae coming denser, 0.5-1 PD apart, dorsally with dorsal spot, femora with ventral sur- and laterally; area between eye margin and face yellow; metasomal terga I-IV with lat- antennal socket with dense small punctures, eral spot; wing membrane lightly brown- contiguous to 0.5 PD apart; antennal lobes tinted, darkest in medial cell, veins dark evenly rounded; frons punctures larger than brown, stigma yellowish brown medially. on lower face, 1-2 PD apart; vertex sparse- Vestiture: —Long, sparse, silvery. ly punctate, punctures small, 1-4 PD apart; Female. unkno—wn. hindocellus 3 ocellar diameters from eye Type material. Holotype S: Caqueta margin; mandible with long apical tooth. Parque Nacional Natural, Cordillera de los VOLUME 107, NUMBER 3 583 24. intermedius 20. osorioi 21.paradoxus 22.pecki 27). turbulentus 25. desertus 33. intermedius 34. osorioi 35. paradoxus 36.pecki 37. pecki Figs. 14-37. 14-23, Side view of male face, antenna removed. 24. 25. 30. 32. 37. Side \ieu of female mesosoma. 26-28, Apex ofhypopygium, dorsal view. 31, Apex ofhypopygium. side view. 29. 33-36. Posterior view offemale metasomal segment VI. — 584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON Pichachos, 2°44N 74°53W, 1,560 m. Mal- mere slender, elongate and tapering apically aise trap, E. Gonzalez, 1-7 Nov. 1997, No. in lateral view, broadened and slightly 662 (BOGOTA). Paratypes: 4 6, same data hooked apically in dorsal view; aedeagus as holotype (B—OGOTA, DAVIS). without dorsal lobe, one-half or more as Etymology. This species is named bi- long as paramere. Color: Black, with yel- lobatus in reference to the narrowly bilo- low markings; clypeus yellow, except api- bate clypeus. — comedial spot and around tentorial pits, in- Discussion. The most distinctive fea- ner eye margin with yellow stripe extending tures of male bilobatus are the black and to hindocellus, antennal lobe yellow, man- pale yellow coloration, clypeus with the dible yellow medially, posterior eye margin medial projection apicomedially narrowly with yellow stripe along lower three- bilobate and laterally convex to lobate, and fourths, vertex with transverse yellow the forewing marginal cell darker than the stripe; pronotum with transverse medial rest of the wing. The male clypeus appears yellow stripe, interrupted medially, and flat in profile. large lateral yellow spot, scutum with pos- teromedial yellow spot and yellow lateral Merithynnus bogotanus (Turner) stripe, scutellum with two large medial and (Figs. 5, 16, 27, 40, 50) lateral yellow spots, metanotum yellow me- Elaphroptera bogotana Turner 1910:196. dially and laterally, propodeum with large Syntype males: Colombia: Bogota (LON- U-shaped yellow sublateral spot; forecoxa DON, BERLIN). ventrally yellow, mid and hindcoxae with — dorsal and ventral yellow spot; femora with Male. Body length 13-15 mm. Head yellow ventral stripe, rest oflegs black; me- (Figs. 5, 16): Cypeus nearly impunctate, tasomal tergum I with large, irregular U- strongly produced apicomedially, apex shaped yellow spot, segments II-IV with sharply bidentate; area between antennal large lateral tergal and sternal yellow spots, 0s.o5c—ke1t.0anPdDeyaepamratr;gifnrownisthansdmalvlerptuenxctwuirtehs tergum V with large lateral yellow spot; punctures 0.5 PD apart; hindocellus three wing membrane faintly brown, veins brown. Vestiture: Silvery. hindocellar diameters from eye margin; fla- — Female. unknown. gellomere I 2.5-2.6X as long as broad; fla- — gellomere II 3X as long as broad; flagel- Material examined. Colombia: Bogota lomere XI 5.0-5.3X as long as broad; man- (2 syntype males). Discussion. Merithynnus bogotanus dible medially and subbasally angulate, broadest submedially, with long acute api- and turbulentus are similar in coloration cal tooth, and smaller subsidiary tooth. Me- and overall appearance in the males. Male sosoma: Pronotum polished, nearly im- bogotanus can be distinguished by the punctate; scutum polished, punctures 1 PD shape of the clypeus, and the more exten- apart; scutellum impunctate medially, punc- sive yellow markings. The male clypeus has tures 1—2 PD apart laterally and anteriorly; the apicomedial margin obtuse or rounded, metanotum impunctate; propodeum with and the face has a complete yellow stripe small punctures 3-4 PD apart, nearly im- that extends along the inner eye margin punctate above petiolar socket; mesopleu- from the clypeal margin to beyond the hin- ron polished, punctures 1-2 PD apart. Me- docellus in bogotanus. In turbulentus the tasoma: Integument densely, finely sha- clypeus apicomedial margin is straight and greened, punctures shallow, 1-4 PD apart; the yellow stripe along the innereye margin hypopygium apically broadly rounded, with barely extends to the ocelli. In addition, the dorsal surface irregularly wrinkled (Fig. marginal cell in bogotanus is only slightly 27). Genital capsule (Figs. 40, 50): Para- darker brown than the rest of the wing; VOLUME 107, NUMBER 3 585 43. nasalis 44. osorioi 45. paradoxus 46. pecki 47^ turbulentus 48. acuminatus 49. bilobatus 50. bogotaensis 51, mimulus 52. montanus 53. nasalis 54. o5or/o, 55. paracfa«« 56. /,er^/ .^ ,„r/,«/em.^ Figs. 38-57. 38-47, Male genital capsule; side view (left). 48-57, Dorsal view (right).