J. HYM. RES. Vol. 3, 1994, pp. 119-132

A Review of the Agile Species Group of (: Sphecidae: )

Alexander V. Antropov

Zoological Museum of the Moscow State University, Herzen Street 6, Moscow K-9, 103009 Russia

— is are a is for Abstract. The agile species group of Pison redescribed, the 12 included species reviewed, and key provided Palearctic is identification. The agile group is restricted to the Oriental and eastern Regions, although one species, koreense, adventive in North America. Five species are new: agiloides from Sri Lanka; chrysoptilum from Borneo; mngyuenfuense from southwestern China; vechti from Malaya and Indonesia; and pulawskii from India. Other species of the group are: agile (Smith) from southern India and Sri Lanka; erythropus Kohl from western India; koreense (Radoszkowski) from eastern Asia and North America; rothneyi Cameron from southeast Asia; browni (Ashmead) from the Philippines; different Turner from Assam, India; and hissancum Gussakovskij from Uzbekistan and Tajikistan. Lectotypes are designated for agile, differens and rufipes (Smith) and a neotype is designated for koreense. Pison koreense is removed from synonymy with agile.

INTRODUCTION CAS California Academy of Sciences, San Francisco,

California (W. J. Pulawski). The Pison, which contains nearly 200 NMNH Nationaal Natuurhistorisch Museum, Leiden, The Netherlands van described species (Bohart and Menke 1976; Menke (C. Achterberg). OUM Hope Entomological Collections, University Mu- 1988), is well represented in all faunal regions seum, Oxford, England (C. OToole). except North America where only an adventive USNM U.S. National Museum, Washington, DC, USA (K. is known. species, koreense, V. Krombein, A. S. Menke). In this paper I review the agile species group ZIN Zoological Institute, Russian Academy of Sciences,

St. Russia I. which contains 12 species, five of which are new. Petersburg, (V. Tobias). ZMK Museum, Denmark Members of this assemblage have been placed in Zoological Copenhagen, (O. Lomholdt). the subgenera Pisonoides and Krombeiniellum, but Menke (1988) used species groups rather than The specimens from the collection of the Zoo- subgenera for infrageneric groups. The agile group Museum of the Moscow State is restricted to the eastern Palearctic and the Orien- logical University (ZMUM) were also used in the review. tal Regions, except for the east Asian species koreerxse which was introduced into North America after World War II (Krombein 1958a). presumably THE AGILE GROUP Morphological terminology used here follows Bohart and Menke and Menke The (1976) (1988). This group is characterized as follows: com- abbreviations are used in the text: following OOD pound eyes densely setose, antenna clavate, oc- = ocello-ocular distance; OD = ocellus diameter; cipital carina complete or nearly so, anterior POD = distance between ocelli. posterior pronotal pit small, subomaulus present, episternal The museum and institutions lent following sulcus straight, forewing with only two submar- for this are used specimens study (abbreviations ginal cells (true second submarginal cell lost in the text): through diminution), hindcoxa without dorsolat- eral carina, lamelli- AUZM Universiteit van Amsterdam, Zoologisch Museum, metapleural flange usually Amsterdam, The Netherlands (W. Hogenes). form, propodeum without lateral carina or crenu- BMNH The Natural History Museum, London, England late ridge, male sternum VIII narrow, gonostyle (C. R. L. Ficken). Vardy, simple, volsella small, penis valve compressed 120 Journal of Hymenoptera Research

laterally and without teeth or notched ventrally. (really 3rd) usually not petiolate; recurrent veins Menke (1988) regarded the setose eyes, the received by 1st and 2nd submarginal cells or 2nd clavate antenna, the two submarginal cells, the recurrent vein interstitial between 1st and 2nd presence of a subomaulus, the straight episternal submarginal cells; hind coxa dorsum with low sulcus, the lamelliform metapleural flange and inner carina, without outer carina; legs finely sculp- claw shape as apomorphies of the agile group. Its tured, without stout spines on tibiae and species apparently represent a monophyletic as- tarsomeres; all tarsomeres IV with small plantulae; semblage. Morphologically they are very similar tarsal claw thick to just before apex; propodeum and differ mainly in leg color, tergal bands, punc- rounded, without lateral carinae or lines of foveae tation, vestiture and proportions of tergum I. Male and crenulate ridges, punctate with smooth genitalia are also similar. One of the new species, interspaces; propodeal dorsum not delimited pulaivskii, stands apart from the others in the group (browni with shallow lateral sulcus), with medium because its metapleural flange is narrow. I regard furrow containing short to complete ridge; abdo- it as a plesiomorphic state. Two other species have men compact; tergum I simple or with preapical in unique autapomorphies: browni the propodeal transverse depression (Figs. 15-19); apical bands dorsum is delimited a laterally by shallow, broad of terga often translucent and with silvery or golden sulcus and tergum I has a distinct preapical de- pubescence; male sternum VIII long, narrow, in pression; agiloides submarginal cell II is open rounded or weakly notched apically; genitalia distally. compact, compressed laterally; volsellae small, —Inner Description. orbits of eyes moderately rounded and weakly setose; gonostyle triangular, emarginate, parallel (eyes equidistant at vertex simple, with long, coarse lateral setae curved be- and or clypeus) slightly converging below (rarely neath; penis valve compressed laterally, consider- above); eyes covered densely with short, erect ably widened apically, without teeth or notches setae (Fig. 3); clypeus convex, in female rounded ventrally. in male or Included — (Figs. 4a, 5a), angulate prominent apically Species. agile (Smith), agiloides sp. n., frons antennae (Figs. 4b, 5b); convex; clavate, com- browni (Ashmead), chrysoptilum sp. n., differens paratively short, with distal flagellomeres wider Turner, erythropus Kohl, hissaricum Gussakovskij, than labrum truncate or long; subquadrate, slightly koreense (Radoszkowski), ningxjuenfuense sp. n., carina a emarginate apically; occipital complete pulaivskii sp. n., rothneyi (Cameron), vechti sp. n. almost — (or complete) circle, narrowly separated Biology. Information is available for two spe- from hypostomal carina; male mandible simple, cies of the group, koreense and erythropus. The that of female with inner tooth slightly distad of former was studied by Iwata (1964) in Japan and midpoint; pronotum with small round pit anteri- by Sheldon (1968) in North America. The Indian without orly, lamellae; scutum and mesopleuron species, en/thropus, was studied by Home (1870). moderately, uniformly punctate; episternal sul- These construct small, clay cells, placing cus almost straight, not curved forward ventrad; them separately or in groups on variable surfaces, subomaulus recurved ventrad; omaulus and ac- but not within linear cavities (borings in wood or etabular carina sulcus absent; mesopleural paral- empty stems of plants). The groups may include leled anteriorly by a row of foveolae; metapleuron up to 21 cells which do not merge in a common smooth; metapleural flange usually broadly lamel- mass but keep their independence. Prey consists liform of posteriorly (Figs. 6-8); tegula entirely punc- 6-31 paralyzed, small, usually immature spi- tate; forewing media diverging after cu-a; forew- ders. ing with two submarginal cells (Figs. 10-13), 2nd

KEY TO SPECIES OF THE AGILE GROUP

—1 All femora completely reddish 2 Mid- and hindfemora brown, forefemora mainly brown 6 narrow India 2(1)— Metapleural flange (Fig. 9); pulaivskii sp. n. Metapleural flange broadly lamelliform posteriorly (Figs. 6-8) 3 Volume 3. 1994 121

3(2) Pronotum, scutum, scutellum, metanotum and propodeal dorsum and hindface with suberect golden setae; Borneo chrysoptilum sp. n. Thorax and propodeum with only silvery pubescence 4 4(3) All legs including tarsi and base of trochanters yellowish-red; western India erythropus Kohl — Trochanters, hintibiae apically and tarsi dark brown 5 5(4) Abdominal tergum I densely punctate (punctures separated by a diameter or less), dull due to dense microsculpture; translucent apical bands of terga I-III whitish, at middle hardly broader than hindtarsal — diameter; Uzbekistan, Tajikistan hissaricum Gussakovskij Abdominal tergum I finely, sparsely punctate (punctures more than a diameter apart), surface smooth, weakly shiny in spite of microstriae; translucent apical bands of terga golden, broad, those of 1 1— 1 1 1 at of middle almost equal to diameter hindtibia; Malaysia, Indonesia vechti sp. n.

Abdominal I same as —6(1) tergum polished, sculpture nearly following terga 7 Abdominal tergum I dull or weakly shiny in contrast to following terga 8 7(6) Tibiae reddish; scutal punctures fine, dense, less than a diameter apart; translucent apical bands of abdominal terga golden, apical bands of terga I-III at middle at least twice as broad as diameter of hindtarsomere cell II Sri Lanka n. — I; submarginal open distally (Fig. 14); agiloides sp. Mid- and hindtibiae brown, foretibia partly reddish; scutal punctures coarse, more than a diameter apart; translucent apical bands of abdominal terga mainly whitish, those on I-III equal to or hardly broader than diameter of hindtarsomere I (Fig. 15); submarginal cell II closed; southern India and Sri Lanka agile (Smith) 8(6) Lamelliform part of metapleural flange mainly dark; translucent apical bands of abdominal terga not broader than diameter of hindtarsomere I 9

Lamelliform part of metapleural flange reddish at least posteriorly; translucent apical bands of abdominal terga (at least of terga II-III) obviously broader than diameter of hindtarsomere I 10 9(8) Abdominal tergum I dull, very densely, finely sculptured, transversely depressed preapically (Fig. 17); apical bands of abdominal terga II-III dark brown, that of tergum I narrower than hindtarsal diameter; propodeal dorsum delimited by shallow depression at least laterally; Philippines — browni (Ashmead) Abdominal tergum I shiny dorsally, coarsely, sparsely punctate, faintly transversely depressed preapically; apical bands of abdominal terga II-III yellowish, band on tergum I almost as broad as hindtarsal diameter and half as wide as band on tergum II; propodeal dorsum not delimited by depressions; southeastern China ningyuenfuense sp. n. - 10(8) Translucent apical bands of abdominal terga I 1 1 bright golden, of equal width; propodeum as long as wide (seen from above), with hind surface mostly punctate apically; ocelli (especially in male) small (OOD>OD); India: Assam differens Turner Translucent apical bands of abdominal tergum I very narrow or absent; propodeum wider than long (seen from above), hind surface transversely carinate apically; ocelli larger (OOD

female I often II-III tergum absent, narrow on terga (hardly broader than diameter of hindtarsomere I), bright golden; tergum I dull due to dense microsculpture; metapleural flange with narrower, densely pubescent lamella (Fig. 11); Russian Far East, Korea, eastern China, Japan, United States koreense (Radoszkowski) — Abdomen II-III 1 .5 comparatively longer, terga times as wide as long; translucent apical band of tergum

I those on II-III I present, terga whitish-yellow; tergum weakly shiny, with sparse microsculpture; with metapleural flange broader, spoon-shaped, sparsely pubescent lamella (Fig. 8); southeast Asia, Malaysia, Indonesia rothneyi Cameron

P. Bohart Pison agile (Smith) (Pison) agile: and Menke 1976:333 (listed). ^ Menke 1988:38 of Figs .3 4 6 10 15 agile. (member agile group, redescnbed).

Parapison agilis Smith 1869:300. Lectotype: female, "Ceylon" Lectotype Selection.—The natural History Mu- Sri , . . (now Lanka) (BMNH), present designation. , f_ ... ~ ... , , ... SeUm haS tW0 femaleS Wlth Smith S handwritten P. (Paraxon) agile: Kohl 1885:186 (listed). e labels. The first female has the museum white P (Ptsonoides) agilis: Turner 1916:616 (new combination, re- tyP described). round label (1) with "Type" printed inside a red 122 Journal of Hymenoptera Research

koreense

MAP 2 ^^ c?

of the Pison in the Old World. Fig. 1. Geographic distribution agile species group P. in the World. Fig. 2. Geographic distribution of koreense New Volume 3, 1 994

Figs. 3-19. Morphological features of Pison species. 3, head in frontal view of P. agile (a, female; b, male). 4-5, clypeus in frontal view P. P. (a, female; b, male): 4, agile; 5, pulawskii. 6-9, right metapleuron in lateral view (arrow shows metapleural flange): P. P. P. 6, agile; 7, koreense; 8, rothneyi; 9, P. pulawskii. 10-13, submarginal cell area of right forewing: 10, P. agile; 11, P. browni; P. P. 12, vechti; 13, pulawskii. 14, submarginal cell area of forewing of P. agiloides (a, right; b, left). 15-19, abdominal tergite I (a, lateral view; b, dorsal view): 15, P. agile; 16, P. koreense; 17, P. browni; 18, P. t'fdifi; 19, P. pulawskii. 124 Journal of Hymenoptera Research

—This is the darkest in the oval, a pale-blue round label (2) with "India" Discussion. species from other members the handwritten on the upper side and "56/150" hand- agile group, differing by written in two lines on the under side, a white narrow, brown apical bands on terga II-III, by the transverse of rectangular label (3) with "agilis Sm. Type" hand- strong, preapical depression tergum written in two lines, and the museum white rect- I and the shallow depressions that delimit the dorsum. The last character is in angular label (4) with "B.M. TYPE. HYM." printed propodeal unique in two lines and "21.538" handwritten below. the group. — Another female has a pale-blue round label (1) Range. Philippines. — with "Ceylon" handwritten on the upper side and Material Examined. 1 male, 2 females. "61 /36" on the under side, a pale-blue rectangular LUZON: Manila (USNM), Laguna (CAS). Tsuneki recorded the from label (2) with "P. agilis Smith. Type" handwritten (1983) species Bontoc, Luzon; in two lines. Only "Ceylon" was mentioned as the Mambucal, Negros; Cagayan de Oro, Mindanao. in the so the type locality original description Pison differens Turner "India" type may not be a true type. The Natural P. (Pisonoides) Turner 1916:617. female, History Museum also has more than a dozen agile differens Lectotype: India, Assam" (BMNH No. 21.540), females from "Ceylon" and three of them have the "Shillong, present designation. same round labels found on the second pale-blue P. (KrombeinieUum) differens: Bohart and Menke 1976:337 (new I believe them to be the members of the "type." combination, listed).

I the with P. 1988:38 of type series and have selected specimen differens: Menke (member agile group). Smith's type label as lectotype and the three other Lectohfpe Selection.—Turner described this spe- females as paralectotypes. I am discounting the cies from three females without selecting a holo- Indian as a specimen type. have two identical labels: — type. They rectangular Discussion. Pison agile is a member of a sub- (1) "Shillong. 5.03" handwritten in two lines and of species with mainly dark legs. Together group (2) "Assam. R.Turner. 1905-125" printed in three with n., it differs from other such agiloides sp. lines ("Shillong, Assam, 5000 ft. (Turner), May" in fine species having comparatively was mentioned in the original description). One microsculpture of abdominal tergum I in which female has in addition Turner's label "Pison are The the interspaces polished. remaining terga (Pisonoides) differens. Turn. Type." handwritten are similarly polished. Pison agile differs from in four lines and two museum labels, a round one and agiloides in having completely brown mid with "Type. H.T." printed in two lines inside a red hind legs, sparser and coarser scutal punctures ring and a white rectangular one with "B.M. and narrower whitish translucent apical tergal TYPE.HYM." printed in two lines and "21.540" bands. handwritten below. I have selected the last female —Known from southern India and as and the two others as Range. only lectotype — paralectotypes. Sri Lanka. Discussion. Pison differens is similar to rothneyi Material Examined.—2 males, 18 females. IN- in color of the body and of the translucent apical bands. The female differs from DIA: Anamalai Hills, Cinchona (BMNH, NMNH). tergal rothneyi by its small lateral ocelli and SRI LANKA: North Western Prov., Kurunegala comparatively elongate with its hindface Dist, Kurunegala; Central Prov., Kandy Dist., propodeum punctate (trans- carinate in The male also has Kandy, Udawattakele (BMNH, USNM, ZMUM). versely rothneyi). very small ocelli but the propodeum is not elon- In both sexes the dorsum is Pison browni (Ashmead) gate. propodeal pol- ished, finely sparsely punctate. The scutum is Figs. 11, 17 finely densely punctate in females. I think that Pisonoides browni Ashmead 1905:961. Holotype: male, Philip- differens may be a local form of rothneyi but more "Manila" no. examined. pines, (USNM 8332), material will be required to resolve this. P. (Pisonoides) browni: Turner 1916:617 (new combination, Range. —Known only from Assam State, In- listed). dia. P. (KrombeinieUum) browni: Bohart and Menke 1976:337 (new Material Examined.—1 3 females. IN- combination, listed). male, P. (KrombeinieUum) browni: Tsuneki 1983:81 (redescribed) DIA: Assam State, Shillong, IV. 1903, V.1903 P. browni: Menke 1988:38 (member of agile group). (BMNH). Volume 3, 1 994 125

Pison erythropus Kohl Pison hissaricum Gussakovskij

Smith 1869:299. "India" Parapison rufipes Lectotype: female, P. (Parapison) hissaricum Gussakovskij 1937:622. Holotype: (BMNH No. 21.539), present designation. Nee Pisomtus female, Tajikistan, "Stalinbad" (now Dushanbe) (ZIN), rufipes Shuckard 1838:79 (now in Pison). examined Parapison rufipes Home, in Home and Smith 1870:165 (biol- P. (Pison) hissaricum: Bohart and Menke 1976:336 (new com- ogy)- bination, listed). P. in and Smith 1870:188 P. hissaricum: rufipes Smith, Home (redesenbed). Menke 1988:38 (member of agile group). Pison (Parapison) erythropus Kohl 1885:183 (new name for Parapison rufipes Smith 1869, nee Ptsonitus rufipes — Discussion. Pison hissaricum also belongs to Shuckard 1838). the subgroup whose members have col- P. (Parapison) erythropus Kohl 1885:186 (listed). brightly ored This can be the P. erythropus: Bingham 1897:221 (listed). legs. species recognized by P. Pisonoides) erythropus: Turner 1916:616 (listed). following combination of features: legs except P. (Pison) Bohart and Menke 1976:335 (new com- erythropus: trochanters, tarsi and tibiae reddish apically; ter- bination, listed). gum I dull due to dense microsculpture of P. erythropus: Menke 1988:38 (member of agile group). interspaces, punctures of tergum I dense; and whitish translucent bands of I-III nar- Selection. —The Natural apical terga Lectotype History row. Museum has three females of this species bearing —Known from Uzbekistan and round labels with "India" handwritten on the Range. Tajikistan. upper side and "98.69" on the under side. One of Material Examined. —8 males, 10 females. them also has a rectangular label with "rufipes Sm. UZBEKISTAN: Aman-Kutan (ZMUM). TAJI- Type" handwritten in two lines. I believe them to KISTAN: Dushanbe; Varzob Valley, Kondara (ZIN, be the members of the series and I have type ZMUM). selected the last female as lectotype and the other two females as paralectotypes. — Pison koreense (Radoszkowski) Discussion. Pison erythropus belongs to the Figs. 7, 16 subgroup whose members have brightly colored legs. This species is easily recognized by its en- Paraceramius koreensis Radoszkowski 1887:433. Holotype: fe- tirely reddish-orange legs (including tarsi and male, "Koree" (Krakow, Poland. Lost). Neotype: fe- basal of the parts — trochanters). male, Russia, Primorskiy Kray (ZMUM), present desig- Biology. The nests observed by Home (1870) nation. in northwestern Ceramius koreensis: Morawitz in Dalla Torre 1894:3 (new India as Parapison rufipes (Sm.) combination, listed in Vespidae). included "a mass of loosely arranged cells of earth Pison (Parapison) koreense: Kohl in Dalla Torre 1897:712 (new attached to some such as a hanging objects, creeper, combination, listed in ). or tendril, pendent straw, or even a curled dry Pison (Pisonoides) koreensis: Turner 1916:617 (new combina- leaf." Home mentioned that the cells were "very tion, listed). P. (Pisonoides) koreensis: Yasumatsu globular" and that their walls consisted of com- 1935:229 (listed). P. (Parapison) koreense: Gussakovskij 1937:622 (listed). paratively large "pellets. ..loosely attached to one P. (Pisonoides) koreensis: Yasumatsu 1939:83 (listed). another." He also "counted ("smallest eighteen P. (Paraceramius) koreense Krombein 1958a:166 (first record in in two " Home) chambers." Nothing from United States). was reported about the number of cells per nest, P. (Paraceramius) koreense: Krombein 1958b:189 (listed). KrombemieUum koreense: Richards 1962:118 name for the species or even family of the prey nor was the (new cocoon described. Paraceramius Radoszkowski 1887, nee Saussure 1854). P. (Pisonoides) koreensis: Iwata 1964:1 —Known with from west- (biology). Range. certainty only P. (KrombemieUum) koreense: Krombein 1967:394 (new combi- ern India. nation, listed). — P. Material Examined. 2 males, 9 females. IN- (Krombemiellum) koreense: Menke 1968a:7 (listed). P. (KrombemieUum) koreense: Menke 1968b: 1102 DIA (BMNH, ZMUM); Maharashtra District, West- (redesenbed). P. koreense: Sheldon 1968:107 em Ghats, Lonavale (NMNH). (biology, larval morphology). P. (KrombemieUum) koreense: Hawkins 1974:279 (biology). P. (KrombemieUum) koreense: Bohart and Menke 1976:337 (listed). P. (KrombemieUum) agile: Krombein, et al 1979:1641 (synony- mized). 126 Journal of Hymenoptera Research

P. koreense: Kazenas 1980:92 (listed). plants, as in erythropns, is unknown. The cells were P. agile: Menke 1988:38 (listed). provisioned with six immature spiders of the ge- — nus Aranens or Neototype Selection. At my request, A. P. (Araneidae) (Iwata 1964) by 20-31 small of the Rasnitsyn recently searched for the type oikoreense spiders genus Dictyna (Dictynidae), in Radoszkowski's collection in Krakow, Poland including males and females of D. bellans Chamberlin and D. sublata Hentz but could not find it. I presume it was lost and I (Sheldon 1968). P. formise cocoons are have selected a specimen from the southern part of cylindrical (6.0-9.1 X 2.0-3.6 to Primorskiy Kray (32 km SE Ussurijsk, 25. VIII. 1986; mm) and, according Sheldon, were surrounded "delicate silken threads eastern Russia not far from Korea)as the neotype. by attaching cocoon to inside of Discussion. —Pison koreense is the best known cell." Sheldon also mentioned a large number of adult Melittobia member of the agile group because its biology has chalybii Ashmead been studied both in its native habitat in Japan (Eulophidae) which were reared from koreense (Iwata 1964) and in the United States (Sheldon cocoon "from a nest constructed in an unsealed cell of 1968) where it was introduced apparently after politum." World War II (Krombein 1958a). I also observed nesting and hunting activities Krombein (1979) synonymized koreense with of koreense during 1983-1986 in the Primorskiy in the Far East of Russia. The agile and Menke (1988) accepted this synonymy. Kray females made their small cells in the After comparing the syntypes of agile with mate- clay between logs wall of an old rural rial from Sri Lanka, Japan, Russian Far East and shed, placing them in groups of up to 20 the United States, I have concluded that koreense is independent cells. Five to seven specimens of a It immature of the Araneus distinct species. differs from agile in having a genus were put into each cell. dull tergum I that contrasts with a shiny tergum II When hunting the female of koreense I looked for the on their webs (tergum shiny in agile). Pison koreense also differs prey and attacked them from the from agile in having bright golden translucent dorsum, embracing the carapace bands on II-III and a to the venter of the apical terga (whitish in agile), largely applying single sting After the the yellowish tibiae (brown in agile) and a transversely spider. paralyzing spider, female turned carinate propodeal hind surface (mostly punctate the prey venter up, clutched the base of its chelicerae with her mandibles it in agile). The metapleural flange of koreense is broad and flew with to and reddish it is in her cell. I observed once and it was (Fig. 7); narrow and dark agile egg-laying only Pison the last before the cell. (Fig. 6). koreense differs from differens and operation sealing Sheldon rothneyi in having a more compact abdomen and a (1968) also mentioned that the egg was attached to the of the weakly developed translucent apical band on ter- abdomen last spider in the cell. gum I. Thus, populations of koreense in Russian Far — the Biology. The wasps were studied in detail in East, Japan and United States have similar Japan by Iwata (1964) and in North America by biological features except prey. Spiders of the Sheldon (1968) who also provided a detailed de- family Dictynidae were preferred in North America and Araneidae in scription of the immature stages of koreense. Both — eastern Asia. Iwata and Sheldon observed that koreense con- Range. Russian Far East, Korea, Japan, east- structed small ern United States: (6.0-10.2 X 4.0-6.0 mm), fragile, China, Illinois, Kansas, Mary- mud cells with land, New Wisconsin. cylindrical finely cemented walls. Michigan, York,— Virginia, The cells are placed separately or in groups of up Material Examined. 28 males, 18 females. to 21 independent cells. The cells are attached in RUSSIA, Primorskiy Kray: Sichote-Alin, Vangou; 32 SE various protected places: inside a photographic km Ussurijsk; Partizansk; 70 km ENE tank (Krombein 1958a); in a culvert, in small de- Partizansk (CAS, USNM, ZMUM). CHINA, pressions and cracks and under a bridge (Sheldon Zhejiang: Hangchow (USNM). JAPAN, Fukui, Mt. 1968); in the nooks of a mud wall under eaves Haku: Ichinose; Chugu; Mie (USNM, ZMUM). (Iwata 1964); and even within the old empty cells USA: Kansas, Lawrence; Virginia, McLean of Trypoxylon (Trypargilmn) politum (Say) (Sheldon (USNM). 1968; Hawkins 1974). Construction of cells on 127 Volume 3. 1994

femora Pison rothneyi Cameron labrum, tibiae except apically, apically, lobe and Fig. 8 pronotal posteriorly, tegula metapleural of flange; translucent apicalbands abdominal terga P. (Parapison) rothneyi Cameron 1897a:81. Holotype: female, I-V and sterna II-V, and terga I-V laterally yellow- India, West Bengal State, "Barrackpore" (now Barakpur) ish-white. (OUM), examined. 1897a:25. male frons P. (Parapison) crassicorne Cameron Holotype: Clypeus, beneath, pronotum anteriorly, West (female in original description), India, Bengal State, scutellum, metanotum and propodeum with dense (now (OUM), examined. Syn- "Barrackpore" Barakpur) erect silvery pubescence, longest setae on onymy by Turner (1916:617). propodeum laterally and posteriorly; pronotal P. (Pisonoides) rothneyi: Turner 1916:617 (new combination, collar, scutum, and abdomen with listed). mesopleuron combi- setae P. (Krombeiniellum) erythropus: Tsuneki 1974:637 (new suberect, dense, short pubescence, longest nation, misidentified). on apical band of tergum I. 1976:336 P. (Pison) rothneyi: Bohart and Menke (listed). inner Labrum slightly emarginate apically, of P. rothneyi: Menke 1988:38 (member agile group). orbits almost parallel, OOD

. a carinate apically Also, rothneyi has more coarsely closed by sulcus or carina; metapleural flange sculptured scutum and deeper impressed broad, spoon-shaped, translucent, with long sil- differs from koreense parapsidal lines. Pison rothneyi very setae. abdominal with left forew- by the comparatively longer terga Right forewing (Fig. 14a) with two, translucent bands three outer well developed whitish-yellow ing (Fig. 14b) with submarginal cells, I. Pison also recur- on all terga, including tergum rothneyi submarginal cell of forewings open distally; has a broader, spoon-shaped reddish-orange rent vein II ending on tiny submarginal cell II (left metapleural flange (Fig. 8) and its tergum I is shiny forewing) and interstitial on right forewing. (semidull in differens and koreense). Tsuneki (1974) Abdomen simple; terga shiny, densely punc- misidentified specimens of rothneyi as erythropus. tate, diameter of punctures decreasing from basal tarsi of his con- I The dark femora and specimens to apical terga; tergum with microsculpture, firm this. other terga more polished between punctures; — distributed Range. This is the most widely sternum I similar to frons, other sterna densely, of the It is recorded of Oriental member agile group. finely punctate, shiny; translucent apical bands from eastern India, Thailand, Viet Nam, Malaysia abdominal segments 1 .5 to twice as broad as diam- and Indonesia. eter of first hind tarsomere. Material Examined.— 1 male, 13 females. IN- Male.— Unknown. DIA, West Bengal: Barrackpore (OUM). THAI- Discussion.—The new species differs from ag- LAND, Chieng Mai Province: Fang Horticultural ile in having the dense, fine punctation of the Exp. Station; Doi Inthanon N. P.: Huai Sai Luang scutum and abdomen, the longer propodeum, the (ZMK). Malaysia: Teluk Merban (ZMK); Kuala comparatively short setae of the thorax and abdo- de Lumpur (BMNH). VIET NAM: Tonkin, Poste men, the broad translucent apical bands of the Dong-Dang (ZMUM). INDONESIA, Sumatra: abdominal segments and the reddish tibiae and are Pakanbaru, Solok; Java: Bogor, Semarang metapleural flange. The last features shared reddish (NMNH). with rothneyi, but the latter has completely tibiae, tergum I is shiny and the propodeum is Pison new scutal agiloides Antropov, species elongate. Pison differens also has fine, dense 14 its Fig. punctures, but it differs from agiloides by larger OOD, by t its dark-brown tibiae and by the weakly Female.—Black Description of Holotype except shiny surface of tergum I. the following reddish: palpi, mandible largely, The open second submarginal cell of the unique 128 Journal of Hymenoptera Research

individual Recurrent vein I on cell I, specimen of agiloides may prove to be ending submarginal when more material is available. The presence of recurrent vein II ending on submarginal cell II near its base interstitial the true second submarginal cell in the left forew- (nearly ). that 7.6 mm. ing is likely to be atypical also, but it illustrates Length the loss of this cell is achieved via diminution of its Male.—Unknown. Discussion. —Pison differs from size in the agile group (see also Menke 1988). ningyuenfuense similar browni an undefined Etymology.—The ending of the species name the most by having dorsum sulcus or emphasizes the likeness of this species and agile. propodeal (no limiting carina), Range.—Sri Lanka. by the comparatively shallowly depressed apical of which is more Type.— Holotype female: SRI LANKA, part abdominal tergum I, coarsely Sabaragamuwa Province, Ratnapura District, punctate, with interspaces only weakly shiny, and Belihuloya Resthouse, 9.IV.1978, M. D. Hubbard, by the whitish translucent apical bands of terga I- T. Wijesinhe (USNM). V. Etymology. —The species name is derived from new the native name of the Pison ningyuenfuense Antropov, species — holotype locality. Range. Known only from the type locality in — China. Description of Holotype Female. Black except southeastern — female: the following reddish: palpi, mandible largely, Type. Holotype CHINA, Hunan alt. 600- labrum, foretibia anteriorly, midtibia basally, Province, Ningyuenfu, July 24-26-28, hindtibia basally and posteriorly and all spurs; 10,800, D. C. Graham (USNM). apical rim of clypeus, tarsi ventrally and tegula reddish-brown; metapleural flange brown; trans- Pison vechti Antropov, new species 18 lucent apical bands of abdominal terga I-V whit- Figs. 12, ish. Female. —Black Pubescence silvery; clypeus, frons, vertex, Description of Holotype except man- pronotum, scutum posteriorly, scutellum, met- the following bright reddish-orange: palpi, anotum and porpodeum with erect setae, longest dible largely, fore and mid tibiae and fore femur hind femora dorsum on pronotum laterally and on propodeum; gena, completely, mid and except mesopleuron and abdomen with suberect, dense, and hind tibia except posteriorly, tegula and short setae, those on gena and lateral angles of metapleural flange posteriorly, apical band on abdominal tergum I longest; scutum with ex- abdominal tergum I and medial spots on terga II- tremely short, erect setae as on femora. III before translucent apical bands; pronotal lobe Pronotum and scutum finely and densely posteriorly and spical tarsomeres beneath yellow- of I-V sterna punctate, shiny, punctures less than a diameter ish; translucent apicalbands terga and apart; mesopleuron, scutellum and metanotum II- VI bright golden. more sparsely punctate (punctures 1-2 diameters Pubescence of head and thorax silvery, mainly on translu- apart); propodeum coarsely punctate (punctures golden on abdominal terga (especially at least twice as large as those on scutum); abdomi- cent bands laterally); frons, vertex, gena, prono- nal tergum I moderately coarsely, sparsely punc- tum, scutum, scutellum, mesopleuron and met- tate (punctures similar in size to those on anotum with dense, short, erect setae (not longer propodeum), interspaces weakly shiny due to than mid ocellus diameter); propodeum dense microsculpture; sternum I similarly punc- posteriolaterally with longer dense, erect setae tate but shiny; following abdominal segments (almost twice as long as mid ocellus diameter); densely, finely punctate (like scutellum), shiny. abdomen with suberect or almost appressed dense, Translucent apical bands of abdominal terga short setae. II-V as broad as diameter of first hind tarsomere, Labrum truncate apically; median lobe of bands of tergum I and sterna II-V less than diam- clypeus obtusely angulate, with narrow, shiny eter of first hind tarsomere; apex of tergum I apical margin; inner orbits almost parallel; shallowly but distinctly depressed (depressed part OOD

of I v. d. ally as hindtibial diameter (those terga and V Pebaton, 28.VI.1932, J. Vecht; Buitenzorg, II-IV of first v. v. d. and sterna 1.5 times diameter Jnsl. Pin., 15.VI.1929, J. Vecht; Malang, hindtarsomere (Fig. 18). IV. 1933, Betrem; Bogor, 1955, Hamann; Mulie; W.

I cell Z. IV. v. d. Recurrent vein ending on submarginal I, Preanger, Soekaboemi, 1933, J. Vecht; recurrent vein II ending near base of submarginal Ambarawa, Lundeking (NMNH); Malang, IV.1933, cell II, the latter very narrow anteriorly, almost Betrem (ZMUM); Mt. Tijoeng, Djampang Tengah, triangular (Fig. 12). 1.1939, K. M. Walsh (BMNH); Kangean Isl.: Length 7.6 mm. Tembajangan, 11.1936, M. E. Walsh (NMNH). Variation in Females (18 specimens).—Femora and tibiae all bright reddish-orange in two females Pison chrysoptilum Antropov, new species from Buitenzorg, Java (28. VI. 1932) and — Tembajangan, Kangean Island all femora dark Description of Holotype Female. Black except posteriorly in a female from Mulie, Java, and sub- the following reddish-orange: palpi, mandible marginal cell II completely triangular in another except apically, labrum, pronotal lobe posteriorly, female from Buitenzorg (16.X.1941). Length 7.0- tegula mainly, widened part of metapleural flange, 7.8 mm. all femora tibiae — except posteriorly and except Male. As in female except: median lobe of apically; translucent apical bands of abdominal clypeus narrower, acutely prominent apically; terga I-V golden. punctation sparser on scutum (punctures 1-2 di- Pubescence silvery on clypeus, mesopleuron, ameters apart) and mesopleuron (punctures 2-4 propodeum laterally and abdominal sterna, golden diameters apart). Length 5.4 mm. on frons, vertex, gena, pronotum, scutum, scutel- Discussion. —Pison vechti is very similar to lum, metanotum, propodeal corsum and hindface hissaricum, erythropus and differens in having wid- and abdominal terga. Clypeus, pronotal collar, ened metapleural flange and largely bright red- propodeal dorsum and terga with suberect or dish-orange legs. It is easily separated from decumbent dense setae (shortest on abdomen, hissaricum and erythropus in having the broad, longest on pronotum); other areas with erect setae, golded, translucent apical bands of the terga. Fur- shortest on scutum and abdominal sterna, longest thermore, it differs from erythropus in having dark- on metanotum and propodeum posterolaterally. brown trochanters and tarsi and a densely punc- Median clypeal lobe rounded, with narrow tate, weakly shiny scutum. Pison vechti differs semitranslucent brown apical margin; inner orbits from differens in having the larger lateral ocelli and of eyes converging above; OD>POD>OOD. a comparatively short propodeum. The color of Clypeus, frons, vertex, pronotum, scutum, the abdomen of vechti is similar to the rothneyi, but scutellum and mesopleuron finely, densely punc- 130 Journal of Hymenoptera Research tate (punctures 0.5-1.5 diameters apart), shiny; tarsomeres III-V; antennal articles beneath, apical metapleural flange broad, spoon-shaped, widened margin of clypeus, tegula, mid and hind tarsomeres part translucent; propodeum more coarsely, III-V reddish-brown. sparsely punctate laterally and dorsally (at least 2 Pubescence silvery only; clypeus, frons, ver- diameters apart), hindface coarsely and densely tex, pronotal collar, scutum, scutellum, metano- punctate (0.5-1 .0 diameters apart), with transverse tum, metapleuron and propodeum with erect se- ridges ventrallv; propodeal dorsum not enclosed tae, longest on propodeum posterolaterally and by sulcus or carina, with narrow median furrow shortest on scutum (scarsely longer than on containing simple carina; tergum I punctate (l-# femora); gena, mesopleuron and abdomen with diameters apart), shiny in spite of microstriahon; suberect or appressed short setae (longest on api- sternum I with coarser punctures (as on propodeal cal bands of terga and shortest on sterna). hindface), shiny; other abdominal sclerites micro- Labrum truncate apically; clypeus compara- scopically but distinctly, densely and uniformly tively broadly rounded (Fig. 5A); inner orbits of punctate (approximately a diameter apart), shiny; eyes almost parallel; OOD=OD=POD. translucent apical margins of terga I-IV twice as Frons densely punctate, shiny; scutum very broad as diameter of first hind tarsomere (on densely, finely punctate, weakly shiny; tergum V and sterna III-V slightly less than diam- mesopleuron uniformly, sparsely punctate (1.0- eter). 1.5 diameters apart), polished; metapleural flange Recurrent vein I ending on submarginal cell I, not lamellate (Fig. 9); propodeal dorsum obliquely recurrent vein II ending near base of submarginal carinate basally and along median furrow, dor- cell II. sum not enclosed by sulcus or carina, sparselv Length 8.2 mm. — punctate (as on propodeal side) (1-3 diameters Variation in Females (2 specimens). Paratype apart); tergum I uniformly, densely ounctate (as female differs by its smaller size (6.2 mm) and on mesopleuron), shiny; tergum II more densely inner orbits of and almost parallel— eye. finely punctate, shiny; remaining terga weakly Male. Unknown.— shiny because of very dense microscopic punc- Discussion. Unlike all other members of the tures; sternum I dull, punctured as frons; sternum agile group, chrysopdlwn has golden pubescence II shiny, punctate as mesopleuron; other sterna on the frons, vertex, thorax and propodeum. It with dense micropunctures, weakly shiny; trans- resembles vechti because of its brightly colored lucent apical bands of terga I-V and sterna II-IV legs and golden translucent bands on the abdomi- whitish, as broad as diameter of hindtarsomere I nal terga. Pison chrysoptilum and vechti may be two (Fig. 19). forms of one species, but more material will be Recurrent vein I ending on submarginal cell I, necessary to resolve— this. recurrent vein II ending on submarginal cell II Etymology. This species name is derived from near its base (Fig. 13). the Greek words and 7.0 mm. din/sos (=gold) ptilon (=down, Length — fluff) emphasizing— the color of the pubescence. Variation in Females (11 specimens). Mid tarsi Range. Northern Borneo (Sarawak, Brunei). completely reddish-orange in three specimens Types. —Holotype female: MALAYSIA, from holotype locality and one from Sarawak: 4th div., Gn. Mulu, RGS Exp., 17.IX- Kurumbagaram; hind tibiae and tarsi brownish 23.X.1977, D. Hollis (BMNH). Paratype (1 female): posteriorly and antennal flagellum completely BRUNEI: Ulu Temburong, Base camp hut, 300 m, dark-brown in one specimen from Coimbatore. 115 16'E,4 26'N, 16.II-9.III.1982, M. C. Day (BMNH). Length 6.0-7.0 mm. Males (3 specimens).—As in females except: Pison pulawskii Antropov, new species clypeus with acute median prominence spicallv 19 Figs. 5, 9, 13, (Fig. 5b); labrum slightly emarginate; mandibles — without inner teeth. Length 4.8-5.2 mm. Description of Holotype Female. Black except Discussion. —The nonlamellate metapleural the following reddish-orange: palpi, mandible flange of pulawskii sets the species apart from other fore distad of coxal largely, labrum, legs apex, mid in the agile group. Other features of the species are and hind legs except basal parts of coxae and the form of the clypeus, the mostly reddish-orange Volume 3. 1994 131

legs and the narrow whitish translucent apical Hawkins, W. A. 1974. A western record for an introduced bands of the Pison the Kansas abdominal segments. Reddish legs (Hymenoptera, Sphecidae) journal of Entomological 47(2): 279. occur in erythropus and whitish abdominal bands Society Home, C and F. Smith. 1870. Notes on the occur habits of some in erythropus, agile and ningyuenfitense, but hymenopterous from the north-west provinces of these have a lamellate India. With an species metapleural flange. appendix, containing descriptions of — some new Etymology. This species is dedicated to species of Apidae and Vespidae collected by Mr. Home. Transactions the Wojciech J. Pulawski. of Zoological Society of London 7(3): 161-196. Range.—Western and southern India. Iwata. K 1964. notes on — Ethological four Japanese species of Types. Holotype female: INDIA, Rajasthan: Pison (Hymenoptera, Sphecidae). Mushi 38: 1-6. (24 35'N), 27.V.1989, W. J. Pulawski V. L. Uddaipur Kazenas, 1980. Materials to the fauna of digger wasps (CAS). Paratypes (3 males, 9 females): same place, (Hymenoptera, Sphecidae) of the Far East of the USSR. date and the Far East, 80-94. collector as holotype (CAS); same place of of Vladivostok, pp. Kohl, F. F. 1885. Die Gattungen und Arten der Larnden and collector as holotype, 21-25. V. 1989 (CAS, Auctorum. derk. k. Disa Verliandlungen zoologisch-botamschen ZMUM); Gujarat: (=Deesa), 4-6. VI. 1989, W. J. Gesellschaft in Wien 34: 171-268. Pulawski (CAS); Karnataka: 915 K. V. Bangalore, m, Krombein, 1958a. Pison tParaceramius) koreense (Rad.), a 26.V.1980, 30.V.1980, K. D. Ghorpade (ZMK); new adventive in the eastern United States. Enfo- News 69: 166-167. Karikal Territory: Kurumbagaram, III.1947, P. S. mologtcal Nathan (USNM). Krombein, K. V. 1958b. Superfamily Sphecoidea. In: K. et Krombein, V., al., Hymenoptera of America North of Mexico - Synoptic Catalog. Agriculture Monograph 2, ACKNOWLEDGMENTS First Supplement, U. S. Department of Agriculture, Washington, DC. pp. 186-204.

I Krombein, K. V. 1967. In: express my gratitude to all colleagues who lent mate- Superfamily Sphecoidea. Krombein, K. B. D. rial for I V., Burks et al., America North study. am grateful to Dr. A. P. Rasnitsyn for his help Hymenoptera of of Mexico— in searches for the type specimen of koreense in Radoszkowski's Synoptic Catalog. Agriculture Monograph 2, Second U S. of collection. I am extremely grateful to Dr. A. S. Menke who Supplement, Department Agriculture, read earlier DC. 386-421. versions of the manuscript and suggested many Washington, pp. Krombein, K. V., P. D. Hurd, D. R. Smith and B. D. Burks. important improvements of the text and who helped me in 1979. the I thank Dr. K. V. Catalog ofHymenoptera in American North Mexico. English. Finally, Krombein and Dr. W. J. of Pulawski Vol. 2. Smithsonian Institution Press, DC. who critically and attentively reviewed the entire Washington, pp. 1199-2209. manuscript and who provided manv useful comments. A. Menke, S. 1968a. New genera and species of wasps of the tribe from the LITERATURE CITED Trypoxylonini neotropical region (Hy- menoptera: Sphecidae: Larnnae). Los Angeles County Museum Contributions in Science 135: 1-9. W. H. 1905. to Menke, A. S. 1968b. review of Ashmead, Additions the recorded Hymenopter- A the New World species of ous fauna of the Pison. I. The Philippine Islands, with descriptions of subgenus Krombeiniellum (Hymenoptera: new species. Proceedings of the United States National Sphecidae). The Canadian Entomologist 100(10): 1100- Museum 28(1413): 957-971. 1107. C. T. Bingham, 1987. The fauna of British India. Hymenoptera, Menke, A. S, 1988. Pison in the New World: a revision vol. 1. Taylor and Francis, London. 179 pp. (Hymenoptera: Sphecidae. Trypoxylonini). Contribu- Bohart, R. M. and A. S. tions the American Menke. 1976. Sphecid wasps of the of Entomological Institute 24(3): 1-171. World, a generic revision. University of California Press, Radoszkowski, O. 1887. Hymenopteres de Koree. Horae Los + Angeles, London, iv 695 pp. Scoitatis Entomologicae Rossicae 21(3-4): 428-436. P. 1897a. IV. O. W. 1962. Cameron, Hymenoptera Orientalia, or contribu- Richards, A revisional study of the masarid wasps. tions to a of the knowledge Hymenoptera of the Oriental Wm. Clowes and Sons, London. 294 pp. Part V. K. 1968. The zoological region. Memoirs and Proceedings of the Sheldon, J. nesting behavior and larval morphol- Manchester of Pison Literary & Philosophical Society 41(2): 1-144. ogy koreense (Radoszkowski). Psyche 75: 107- Dalla C. G. Torre, 1894. Catalogus Hymenopterorum hucusque 117. descriptorum systematica et synonymicus. Vol. IX. G. En- Shuckard, W. E. 1838. Descriptions of new exotic aculeate 181 Transactions gelmann, Lipsiae. pp. Hymenoptera. of the Entomological Society of Dalla C. G. 1897. London Torre, Catalogus Hymenopterorum hucusque 2(1): 68-82. F. 1869. descriptorum systematica et synonymicus. Vol. VIII. G. Smith, Descriptions of new species of the genus Pison, 750 and a list Engelmann, Lipsiae. pp. synonymic of those previously described. V. V. 1937. Transactions Gussakovskij, Especes palearctictiqques des genres of the Entomological Society of London. 9: 289- Didmeis Wesm., Pison Latr. et Psen Latr. (Hymenoptera, 300. Travaux de ' K. 1974. Sphecodeal). I'Institut Zoohgie de I Academie Tsuneki, A contribution to the Knowledge of des Sciences de I'URSS 4(3-4): 599-695. Sphecidae occuring in Southeast Asia (Hym.). Polskie 132 Journal of Hymenoptera Research

Pismo Entomologiczne 44(3): 585-660. Yasumatsu, K. 1935. The genus Pison Spinola of the Japanese Tsuneki, K. 1983. Further studies on the Larrinae of the Empire (Hymenoptera, Trypoxylonidae). Annotationes Philippine Islands, with remarks on the Indian species zoologicae Japonenses 15(2): 227-238. of the genus Lyroda (Hymenoptera, Sphecidae). Special Yasumatsu, K. 1939. Notes supplementaires sur le genre Publications of the japan Hymenopterists Association 24: 1- Pison Spinola du Japon (Hymenoptera, Trypoxylonidae). 116. Festschrift zum 60 Geburtstage von Professor Dr. Embnk Turner, R. E. 1916. Notes on the wasps of the genus Pison and Strand, Riga 5: 81-84. some allied genera. Proceedings of the Zoological Society of London 1916: 591-629.