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The Classification of Old orldW Anthidiini (Hymenoptera, Megachilidae)

Charles D. Michener University of Kansas

Terry L. Griswold Utah State University

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Recommended Citation Michener, Charles D. and Griswold, Terry L., "The Classification of Old orldW Anthidiini (Hymenoptera, Megachilidae)" (1994). Mi. Paper 169. https://digitalcommons.usu.edu/bee_lab_mi/169

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THE UNIVERSITY OF KANSAS SCIENCE BULLETIN

\bl. 55, No. 9, pp. 299-327 NOV 2 8 19940ctober 20, 1994 HARVARD UNIVERSITY

The Classification of Old World Anthidiini

(Hymenoptera, Megachilidae )

Charles D. MichenerI .\nd Terry L. Griswold-

TABLE OF CONTENTS

Abstract 299 Introduction 300 Acknowledgments 300 List of Genus-Group Names 300 Kev to the Genera of Anthidiini of the Eastern Hemisphere 302 Siipplementan' Kev to Males of ."Viithidiine Genera of the Eastern Hemisphere 304 Comments, Descriptions, and Keys to Subgenera 305 (See page numbers for genera in List of Genus-Group Names) Literature Cited 325

.\BSTRACT

Genus-group names of Anthidiini of the eastern hemisphere are listed with indications of synonymies and status (i.e., genus or subgenus). Keys to genera and stibgenera are provided, together with explanatif)ns of classificatoiT decisions. The taxa are n. tollowing new described: Acanlhidium g., rspe species: Acauthidium balracn. sp. from hidia; Clistnnthid- n. ium, stibg. oi Eoauthidium, type species: Uianlhidium turnmcum Ma\Tomoustakis; Indanthidium n. g., type species: Ind- anthidium n. n. n. crenulaticauda sp. from India; LarinosleUs g., type species: Larinostelis scapulata sp. from Kenya; Trarhiisflidnn. Trachusoides n. g.. tvpe species: simplex sp. from hidia: Tiichanthidindcs. n. siibg. oi Pnrlnnnthidum. rv'pe species: Pachyanthidium scmiluteum Pasteels; '/Mslrranthidium. n. subg. oi Ajrauthidtuw. type species: Nifpymlhidiitm tergofasrialiim Pasteels; and Pseudoaitthtdium hmrli/n/um n. sp. from Tanzania, an untisual species placed in the stibgentis Tuberanthidiuw.

' SM)U En roMOLOGICAL Ml'SEf.M, UNtVTERSITY OF K\N.SAS, LAWRENCE, KWSAS 66045, USA. - Bee Biology & Svstem.\tics Laboratory, Utah State University', Logan, Utah 84322-5310, USA. 300 The UNrvERSIT^ of K\nsas Science Bulletin

INTRODUCTION tain characters, comprehensi\e knowledge of anthidiines was advanced by his contributions. Frustrating as his works can The preparation of an account of bees of the world by be, we must say that a paper such as oius would have been one of us (CDM) led to development of the present paper difficidt to prepare using the scattered literature available on old world anthidiine bees, to serve as a precursor for the before Pasteels" publications. Nonetheless, we believe that account planned for the future. The anthidiine bees con- one of the contributions of the present work is better to in- stitute a tribe in the subfamih' Megachilinae in the sense of dicate relationships among taxa by synon\'mi/ing some of

. Pasteels' names others Roig-j\lsina and Michener ( 1993) We now exclude Dioxyini generic and reducing man)' from from the Anthidiini. The Anthidiini differs from other generic to subgeneric status. tribes (Dioxyini, Megachilini, Osmiini) in the folloiving The new species described herein are all rare, at least in characters: collections; the number of available specimens of each Mandible of female much broader than that of male, varies from one to foiu". Therefore we have gi\en rathei full with three or more teeth, sometimes difficidt to interpret descriptions and illustrations, and have included probable if there is a long and partly smooth apical margin as in some generic characters in the species descriptions, so that ex- neotropical forms. Metanotimi without median tubercle. plicit information will be available on as manv characters Stigma less than twice as long as broad, inner margin basal as possible for the use of those who do not have specimens to vein r usuallv little if any longer than width, rarely about available. 1.5 times width; prestigma commonly short, usually less In the geographical information provided in the keys, than twice as long as broad; claws offemale cleft or with inner Africa means subsaharan Africa and Oriental means the Ori- tooth except in Trachusoides: oiuer surface of hind tibia ental faunal region, i.e., tropical Asia and nearby islands. usually with abundant simple bristles; sting of female well Clearlv a numerical phylogenetic study would have been developed; bodv commonly with yellow or white (some- desirable and will be necessarv before a more definitive tiines red) integumental marks. classification is developed, whether or not that classification We use some terminolog)' that will be explained else- is based strictly on phylogenetic findings. W'e hope that the where in greater detail. The following brief explanations W\\\ present paper, and the world accoimt alluded to above in suffice for this paper: which taxa will be briefly characterized, will at least suggest omalus—the angle between the anterior siuface and the a multitude of characters that might be used in a formal phv- lateral surface of the mesepisternum. logenetic studv. juxtantennal carinae—the pair of more or less longitudi- nal carinae, each just mesal to an antennal base and fre- quently overlapping the antennal base. (Terms like .'SiCKNOWTEDGMENTS interantennal carinae suggest carinae extending between the antennal bases.) We much appreciate the opportunity to examine rvpe specimens and other material lent us b\ the institutions and Tl, SI, etc. —first metasomal tergiun and sternimi, etc. following per- sons: Museum an der Humboldt-Unixersitat, Berlin Thus, Tl is the second abdominal the Zoologisches tergimi, propodeimi (Frank Koch): The Natiual Histon Museum, London (Laraine first. being Ficken); Director, Department of Agriculture, Nicosia, Cyprus; tribe is in all This found continents, biu onlv one species American Museimi of Natural Histoiy, New York (Jerome G. of San Francisco V. is known from Australia. Elsewhere each continent con- Rozen.Jr.) ; California Academy Sciences, (W. Pulawski); University of California, Davis (l.ynn S. Kimsey); South tains many genera and species. There is a tendency for the .\frican Museum. Capetown (V. Whitehead): Zoological Institute, development of numerous, small, morphologicalh' distinc- Russian .Academy of Sciences, St. Petersburg (Y. A, Pesenko); G. tive taxa so that many genera and subgenera ha\e been rec- van der Zanden. Eindlioven. Netherlands. ognized. On the other hand Warncke (1980) placed all CDM's pan in this paper was greatly facilitated by Virginia Ashlock and b\ National Science Foundatit)n DEB-9,S02186. nonparasitic Anthidiini of the western palearctic region in grant the genus Anthidium. This clearlv limips very dissimilar forms into a paraphyletic group from which parasitic taxa must have evolved. LIST OF GENUS-GROUP NAMES In the present paper, we ignore taxa of the western hemi- sphere. Onl\' the following genera occm both in the Amer- Acaulhidium .Michener and Griswold, new 305. icas and in the eastern hemisphere: Anthtdii'Uum, Anthidium, genus; p. Michener; 307. Trachusa, and Stelis. Afmnthidiuni p. Ajraulludinm Michener s. str., 1948: 24. We have not included detailed descriptions except for new BrdnthidiiniiPaateeh, 1969a: 88 = HouiiiithidiumPasteeh, this use of earlier the taxa; paper requires works, especially 1969a: 88. Pasteels 1984). We are major papers by (1969a, impressed Capanthidium Pasteels, 1969a: 85. that, although Pasteels" works showed many signs of haste, Donuinthidium Pasteels, 1969a: 95. he knew a great deal about anthidiine bees. Even though Immanthidium Pasteels, 1969a: 89. his kevs often do not work and his diagrams of structures Mesanthidiellum Pasteels, 1969a: 83. II / i 1950: 316. and his descriptions somedmes conflict or are wrong for cer- Mesa hid am Popov, MCZ . Old World Anthidiixe Bees LIBRARY 301

Pasteels, 1984: 57 = Melantliidium Pas- Pachyanthidium Friese; 317. Nigrauthidium p. JOV 28 1994 teels, 1969a: 90 (preoccupied). Ausanthidiinn Pasteels, 1969a: 60. Omuthidium Pasteels, 1969a: 95. Pachyanthidium Friese s. str., 1905; 66, also described A:«ifl/(//»V/»(/« Pasteels, 1984: 33. as new by Friese, 1908: 15^^Af?\/A F?Pi Zosteranthidium Michener and Griswold. new sub- Trichanthidiodes Michener aWjwfWti^W^iTjcW genus. ffenus. 'VCKoirY Afrostelis CockexeW. 1931: 340; p. 312. Trichaulliidiiiiii Cockerell, 1930: 52. AnthidieUum Cockerell: p. 312. Plesianthidium Cameron; p. 319. Anauthidii'llum Pasteels, 1969a: 49. Carinanthidiiini Pasteels. 1969a: 42.

Aiilhidii'lluin Cockerell s. str., 1904b: 3 = Ccrianthidium Plesianthidiuiii Cameron s. str., 1905: 256. Friese, 1923: 304. Spiiiaiithidii'Uum Pasteels, 1969a: 59. = Chlomuthldit'llii III MdvromousVdk'is, 1963b: 491 C.lilni- Spinaiithidium Mavromoustakis, 1951: 977. anllud/umPa.steels. 1969a: 48 (unjustified emen- Pseudoanthidiiini Friese; p. 319. dation). Exaiithidium Pasteels, 1969a: 82. Clypanthidium Pasteels. 1968: 1060, also described as Miminlhidiuni Cockei-eW, 1930: 45. new in Pasteels, 1969a: 53. Pseudoanthidnim Friese s. str., 1898: 101 = Paranthi- Pyoirnilhidiiim Krombein, 1951: 292 = Pyirnaiithidium diellum Michener, 1948: 25 = ParaunlhidicUum MaNTonioustakis, 1963b: 491 emen- Pasteels, 1969a: 80 emendation) = (imjustiFied (unnecessaiT' dation) = Maxi'omoustakis, Can II I'll Hill Pasteels, 1969a: 80. PygnauthidtrUum' 1963b: 492. Royanthidiuiii Pasteels, 1969a: 86 = Reanthidium Pasteels, RantliidwUum Pasteels, 1969a: 48 = Rhanlhidwlhim Pas- 1969a: 87. teels, 1972: 102 (imjustified emendation). 5>w/crt)v /((//« Pasteels, 1984: 32. Anthidioma Pasteels, 1984: 34; p. 313. Tuberanthidium Pasteels, 1969a: 87. Anthidium Fabricius; p. 313. Rhodanthidiuni Isensee: p. 321. = Anihidium Fabricius s. str., 1804: 364 Mehmoduthid- Asiauthidium Popox, 1950: 315= Trianthidiu»iMa\ro- iiim Tkalcij, 1967: 91 = Echinanlhidhun Pasteels, moustakis, 1958: 435 = Oxyanthidiiim Mavro- 1969a: 101 = Po«to)(//»W»(w Pasteels, 1969a: 105 moustakis, 1963a: 653 = Axillanthidium Pasteels, = Ardenthidnim Pasteels, 1969a: 103 = Morphan- 1969a: 39. thidium Pasteels, 1969b: 423. The last is invalid; Meganthidiuiii Popov, 1950: 315. = no indication of a tvpe species, biu contains RJiodanlhidiiim Isensee s. str., 1927: 374 Bclhiiithidium same species as Ardenthidium. Pasteels, 1969a: 38. = Gulanthidium Pasteels 1969a: 101. SerapiitaCockereW, 1904a: 357 .S>ra/;M Smith, 1854: 218 (pre- Nivanthidium Pasteeh, 1969a: 106 occupied). Pwanlhidium Friese, 1898: 101. Stelis Panzer; p. 322. SeiH'ranthidlum Pasteels, 1969a: 106. Makmthidium Pasteels, 1969a; 26. Turknnlludium Pasteels, 1969a: 103. Protostclis Friese, 1895: 25 = Heterostelis Timberlake, = Apia iithidi II III Piisleeh, 1969a: 41. 1941: 125 Doxanthidium Pasteels, 1969a: 28. 167. A/rr)/;/;(w Pasteels. 1984: 132. Pseitdostelis Popow 1956: Bathaiilhidium .\la\Tomoustakis; p. 314. Stelidomotpha Morawhz. 1875: 131. = Bdthunthidium Mavromoustakis s. str., 1953: 837. 5Yefa Panzer s. str., 1806: 246 Gyrodroma King, 1807: = = Maiithidium Pasteels, 1969a: 43. 198 G)'w??i/iSpinola, 1808; 9 Ceraplastes G\s- Stenanthidiellum Pasteels, 1968: 1059 = ? Lasauthidium tel, 1848; x = Leucostelis Noskiewicz, 1961: 126. Romankova, 1988: 26. Numerous genus-group names proposed for Benanlhis Pasteels, 1969a: 61. Status unknown; p. 314. North .\merican species are also best regarded = Cyphaiithidiiim Pasteels, 1969a: 57 TrianthidicUum Pasteels, as svnonvms of Slelis s. str. 1969a: 58; p. 314. TrachitsaPanzer; p. 323. Eoaiithidiiim Popo\'; p. 314. Archiaiithidium Maxromoustakis, 1939; 91. CJisldiithidiiim Michener and Griswold, new subgenus. CoHgo/rar/n/.s/; Pasteels, 1969a; 24. = Eiuiulliidiiim Popov s. str., 1950: 316 Eoanthidiellum Massanthidium Pasteels. 1969a: 24. Pasteels. 1969a: 51. Metatrachiisa Pasteels, 1969a: 22. Hcmidii'llum Pasleeh. 1972: 112. Orthanthidium Mavromoustakis, 1953; 837. Salemanthidium Pasteels, 1969a: 51. ParaanthidiumFriese, 1898: 101 = ProtanthidiumCock- = = fMCH/iM Gerstaecker, 1857: 460 Z)//o6o/>W/m Fairmaire, 1858; erell and Cockerell, 1901; 49 Protoanthidiuin = = 266 Purnmspis Ritsema, 1874: Ixxi; p. 315. Cameron, 1902: 125 Philotrachusa Pasteels, Gnathdiilhidiiim Pasteels, 1969a: 92 (not Urban, 1992): 1969a; 22. = p. 315. Trachma Panzer s. str., 1804; expl. pi. 14-15 Dijjh- = Irtmiiil/iidiiiiii Michener, 1948: 25. ysis Lepeletier, 1841: 307 Megachilrnides in Indinitliidiiiiii .Michener and Griswold, new genus: p. 315. Radoszkowski, 1874; 132. See Michener, press. Larinostelis Michener and Griswold, new genus; p. 317. Trachusoides Michener and Griswold, new genus; p. 324. Neanlhidium Pasteels, 1969a: 93. 302 The Uni\t,rsiti' of K-vnsas Science Bi'lletin

KEYTO THE GENERA OF ANTHIDIINI OF THE EASTERN HEMISPHERE (See also Siipplementan' Key to Males, below.)

1 . Mandible ol female with 5 to 1 8, usually sharp teeth separated b} acute notches or (in one species of Pach^an- thidinm) minutely denticulate; maxillaiT pal])us short, 2-segmented or in Inddtilhidiuw aj^parentlv l-segmented (arolia absent; base ot propodeal triangle pmictate or finely roughened, nearl) always hair); propodeiun with- out basal series of pits and without fovea behind spiracle; juxtantennal carina absent) 2 —Mandible of female with three or foin- teeth, or if with five to ten, they are roimded and separated b\ rounded emarginations; maxillaiT palpus commonly 3- or 4-segmentecl but sometimes 2-segmented 11 2. T5' with posterior premarginal zone depressed (except sometimes medialh), moie finely pimctate than rest of tergum and usually densely so, not o\er half as wide laterally as mediall)', this zone ending in veiy narrow smooth posterior margin, anterior margin of depressed zone often obtusely angulate medially so that whole zone is very broadlv triangular 3 —T3 with posterior premarginal zone not depressed or weakly to strongly depressed, this zone if recognizable, pimc- tate like rest of tergum to impunctate, often over half as wide laterally as medially, often ending in broad smooth posterior margin, anterior margin of depressed zone straight or cin"ved, not angulate but rarely with small basal median angular projection 4 3. T6 of female with margin usually not denticulate^, sometimes with lateral tooth, with median apical notch or emargination acconnnodating sting; T5 of female with basal edge of depressed marginal zone commonly ob- tusely angulate medially so that zone is broadly triangular; penis valves of male widely separated basally, united by long, narrow bridge; volsella projecting as lobe at apex of gonocoxite (widespread) Authidiiim —T6 of female with margin denticidate, almost always without lateral tooth, usually without median apical notch; T5 of female with basal edge of depressed marginal zone not angulate medially; penis valves close together or fused basally. bridge therefore short or absent; Nolsella small or absent (.-Xirica, southern Palearctic) Afranlhidium (part) 4. First recurrent vein joining first submarginal cell; axilla acutely pointed; face with longitudinal median shiny from frons to —ridge clypeus (Africa) Serapista First recmrent vein entering second submarginal cell; axillar margin roimded or straight; face without longitud- inal median ridge 5 5. Basal area of propodeum hairless except laterally; pale markings absent on body; scutellum not angulate lat- erally; terga without impunctate margins (male unknown) (southern Africa) Anthidioma —Basal area of propodeiun with hairs, sometimes veiy short; pale markings usually present, but if not, as in Gudthaulhidium and Afranlhidium {Immanthidium. etc.). then scutellum usually angulate laterally and terga with impunctate margins 6 6. Preoccipital ridge dorsally and omalus produced as translucent lamellae (Africa) Pachyanthidium (part) —Preoccipital ridge and omalus roimded or at most carinate. except preoccipital ridge lamellate in Gnnthanthidium 7

7. Subantennal suture straight or weakly arcuate; S4 and S5 of male not strongly concave, rather simple, S3 to S5 without combs or areas of specialized bristles, their posterior margins straight or weakly concave (with lateral projections in Neanthidium and on S5 in Gnathanthidium) 8 —Subantennal suture distinctly arcuate outward; S3 to S5 of male usually concave, S4 and S.5 or at least the latter short and largely hidden except in liidanthidiiim, at least S5 with posterior lateral projection except in Pseudoan- thidium (Exanlhidium) and Indaitthidium; S3 often with comb or area of wavy bristles 10 8. T6 and T7 of male each with four large equidistant teeth; S4 and S5 of male with lateral projections; T6 of fe- male with lateral spine and median emargination (length 9-13 mm) (north Africa) Neanthidium —T6 of male simple, T7 short, bidentate, bilobed, or tridentate; S4 and S5 without lateral projections or S."! with such projections in Gnathanthidium:'!^ of female without c^r with veiT weak lateral spine and median emargination

.' 9

9. Mandible of female with 13 or 14 teeth, apex broad, lower two teeth and upper one large, others small and subequal; scutellum transverse, truncate, carinate; tibiae coarsely tuberculate (east Africa) Gnathanthidium —Mandible of female with eight teeth or less; scutellum roimded as seen from abcne, not or incompletely cari- nate; tibiae not tuberculate except in some species of subgenera Gapanthidium and Xcnanlhidium (.\frica, south- ern Palearctic Afranthidium (part)

'' More anterior terga and T6 of males reflect the same features, often less clearly. Taxa that are not clearly .separable by this charac- ter can be run to either alternative.

'*T6 is denticulate in some. e.g.. Anthidium (Proaiitltidnii/ij oblungalum (lUiger). Oil) World /lv/7///)//.\7: Bees 303

10. T7 ol male nearly as wide as T(i, muliidentate; subantennal siiuire arising at upper end of tentorial pit (S3 of male without wa\"\ bristles; S5 with margin strongh concave l:)ut no lateral projections) (southern India) Imldnlhidium —T7 of male markedly narrower than T6, 2- or 3-toothed; subantennal suture arising from epistomal suture well above tentorial pit [except in Psntdnnrilhidium (Rnyanlhidinm) rrtirulntiim Mocsar\] (Palearctic, Oriental, ^\frica) Psi'udnaiitliidiiiiii

1 1. Omalus lamellate, continued onto venter of thorax and there separated Irom middle coxa by less than width of middle trochanter I'J —Omalus lamellate or not, if lamellate often not continued onto \ enter of thorax, but if so, mesepisternum be- tween mid coxa and omalus (howcxer recognized) as wide as or wider than width of middle trochanter 13 11'. Propodemn with fovea defined b\ carina behind spiracle; preoccipital ridge dorsalh roimded or with low ca- rina (Holarctic, Oriental, northern Australia, Africa) Aiithidit'lliini (part) —Propodeum without fovea behind spiracle; preoccipital ridge behind vertex lamellate (.Africa, .southern Asia) Pachyanthidiuni (part) 13. Lower part of preoccipital carina sk)ping forward and continuing directly to lower mandibular articulation; axilla frequentlv pointed posterioih (anterior coxa with lamella in most species; hind trochanter ofniale with

preapical ridge, carina, lamella or tooth on inner surface; arolia absent) (Palearctic, south to Kenya) . . Icternuthidiinii —Lower part of preooccipital carina absent or if present and extending to lower part of head, ending below and mesal to lower mandibular articulation, or if reaching mandibular articulation [in Anlhidiellum (Chloranthidiel- him)]. directed below it and then cur\ing up to articulation; axilla not pointed posteriorly except in some para- sitic genera that lack a scopa 14 1 4. Face with three longitudinal ridges or carinae, twojuxtantennal carinae and median longitudinal one on frons and supraclypeal area that is often only a shiny ridge (body withotU yellow markings) 15 —Face without longitudinal median ridge or carina and usually without juxtantennal carinae 16 15. Mesepistei num in front of middle coxa with strong \'ertical ridge; scopa absent; scutellum produced as two broad, flat lobes oxerhanging metanotum and propodeuiu (Africa, Oriental, eastern Palearctic) Euaspis —Mesepisternum without vertical ridge in front of mid coxa; scopa present; sciuellum rounded and not nuich pro- duced in profile (west Africa) Atwpiuiii l(i. Vein cu-v of hind wing usually half as long as second abscissa of M+Cu or more, oblique; middle tibia as broad as hind tibia or nearK so (T7 of male simple or bilobed) 17 —Vein cu-\' of hind wing less than half as long as second abscissa of M+Cu, oblicjue or transverse; luiddle tibia usu-

ally narrower than hind tibia '. 19

17. Claws of female simple (southern India) . Trarhnsnider. —Claws of female cleft or with inner median or preapical tooth 18 18. T7 of male curled under so that dorsal surface faces downward; mandible of female dull, minutely roughened and with veiy short hairs, carinae absent on basal half of mandible; middle tibia with anterior margin strongly curved so that at lowermost extremity it is usually at right angle to line across distal end of tibia (Holarctic, Africa, Oriental) Tracbusa —T7 of male directed posteriorly although small, short and transverse; mandible of female slightly shining, cari- nae strongly shining; middle tibia with anterior margin less strongly convex, at apex at acute angle to line across apex of tibia (Oriental) Apianthidmm 19. Anterior part of axilla produced to a point or lobe directed laterally, behind which mat gin is concave; T7 of male with median point and two lobes on each side, thus with five a]3ical projections (India) Aanithidium —Axilla rounded or soiuetintes poiiUed posteriorly, or if with basal lateial projection, it is cuned posteriorly; mai- gin of T7 of male with less than five apical projections 20 20. Axilla positioned and produced laterally so that it almost abuts against posterior end of tegula; arolia absent; scopa abseiu (Africa) Lariitostdis —Axilla not abiuting tegula; arolia present except in Eoanthidium (Stdnnaulhidhnn): scopa present except in Stelis and Ajrostelis 21 or that st)me 21. Scopa present; front and middle tibiae each with one apical spine angle except ('^pliaiilhidiiini and Eoanthidium have two spines on middle tibia 22 —Scopa absent; front and middle tibiae each with midapical and posterior apical spine, so that each tibia has two apical sj^ines 27 22. Juxtantennal carinae present although sometimes weak; interantennal distance u.sually less than, rarely equal to, antennocular distance; 86 of female usually with spine or premarginal ridge, sometimes weak and lateral only, so that sternal margin looks thick, sometimes elevated to lateral tooth (T7 of male over half as wide as T6) (Palearc- tic, Africa, Oriental) Eoaiilhidiitin 304 The UNr\T,RsiT\' of Kansas Science Bulletin

—-Jiixtantcnnal carinae completely absent; interantenna! distance iisiiallv greater than antennociilai distance; S6 of female unmodified, with margin thin 23 23. Scntosciitellar suture superficially similar to scutoaxillar siitiue, usually closed but if smooth shining floor of groove is visible, usually not divided into two parts; subantennal suture approximately straight or only slightly ar- cuate; fovea behind propodeal spiracle absent; body usually over 10 mm long, although Cyphanlhidium may be 6.5 mm long 24 —Scutoscutellar suture open to shiny bottom or fovea, thus very different from scutoaxillar suture, shiny area di- vided medially or //suture closed (as in Anthidiellurn s. str.) then subantennal suture strongly arcuate outward; fovea behind propodeal spiracle present, defined posteriorly by carina, but fovea sometimes not larger than spir- acle; body iisualh' 8 mm long or less 26 24. T6 of male with median apical tooth or small projection; bod)' length 8.5 mm or less and metasoma with con- tinuous yellow bands [form and coloration as in Afranthidium (Oranthidhtm)] (southern Africa) Cyphanthidium —T6 of male withotit median apical tooth; body length ustiallv 8.5 mm or more, if less, as in some Plesianthidium or {Spinauthidium}, then metasoma without yellow 25. Yellow or cream markings absent or limited to face of male; T3 and other terga with depressed premarginal zone sublaterally nearly half length of exposed part of tergum; T6 of male with median lobe (often subtruncate and elevated) and lateral tooth so that it is trifid or, in subgenus Spinanthidiellum, trimcate with a longitudinal median ridge at apex (South Africa) Plcsiauthidium —Body with yellow or reddish-yellow markings; T3 and other terga with depressed premarginal zone sublateralh one-third length of exposed part of tergum or less; T6 of male simple or with short, broad, rounded median lobe, sometimes (in RJiodanthidium s. str.) also with lateral tooth and thus trifid (Palearctic) Rliodantludium 26. Omalar carina absent or extending down only to middle of mesepisternum; T7 of male, if trilobed, with me- dian lobe much longer than lateral lobe or spine; subantennal suture straight (eastern Palearctic, Oriental) Bnthanthidium —Omalar carina strong, sometimes lamellate, and extending onto ventral surface of thorax, sometimes across ven- ter except in subgenus Clypanthidium in which omalar carina does not reach lower part of mesepisternum; T7 of male, if trilobed, with median lobe small, either not separated from lateral lobe by emargination or not longer than lateral lobe; subantennal sutme usually arctiate ouUvard (Holarctic, Oriental, northern Australia. .Africa) AnlhuUclhnn (part) 27. Tegula enlarged, especially posteriorly, so that width posteriorly is nearly equal to length; scutum longer than wide (Africa) Afrostelis —Tegula of ordinaiy size and shape, widest medially and not as wide as long: scutum wider than long, only mod- erately so in subgenus Stelidnmorphn (Oriental, Holarctic, south to Kenya) Stelis

SUPPLEMENTARY KEY TO MALES OF ANTHIDIINE GENERA OF THE EASTERN HEMISPHERE

The preceding key will be frustrating for various reasons, but one major reason will be that Couplet 1 is largely based on a character of females: stipplementai"y characters will help, btit as indicated within the couplet, they are not always decisive. The following supplementaiy key for males leads either to certain genera or to couplets in the main key, thus bypassing Couplet 1. In reality, its main fimction is to help identif\' males of taxa that should run to 1 1 in the main key but that lack arolia, as do all taxa that run to 2.

A. Arolia absent B —Arolia present II B. Paleotropical species C —Palearctic species 2 C. Vein cu-v of hind wing more ihan half as long as second abscissa of M+Cu, oblique; middle tibia as broad as hind tibia or nearly so D —Vein cii-y of hind wing less than half as long as second abscissa of M+Cu; oblique or transx erse; mid-tibia nar- rower than hind tibia E D. T7 curled under so that dorsal surface faces downward: middle tibia with anterior margin strongK' cuned so that at lowermost extremity it is usualh at right angle to line across distal end of tibia (Holarctic, Africa, Oriental) Trachusa (part) —T7 directed posteriorly although small, short, and transverse; middle tibia with anterior margin less strongly con- vex, at apex at acute angle to line across apex of tibia (Oriental) Apiantliidiin/i E. Axilla almost entireh lateral to lateral margin of sctUimi; outer, apical margins of fore and mid tibiae each with two minute spines (Kenya) l.ariuostelis Old Wori.d AxTHtDiixE Bees 305 — \xilla at most extending slightly lateral to lateral margin of scutum; outer apical margins of fore and mid tibiae each with at most one spine F F. Omalus carinate for at least three-fourths of distance from upper end to midventral line G —Omalus carinate for no more than half of distance from upper end to mifhentral line 2 G. Preoccipital carina present dorsally, behind vertex H —Preoccipital carina absent dorsally, behind vertex I H. Hind tibia tuberculate on outer surface; sciuelhun \er\- short, width greater than foiu' times length, onlv slightly overhanging metanotimi (for one-third its length) (east .-Vfrica) Gnathunthidinm —Hind tibia not tuberculate; sciuelhmi moderatelv long, width equal to or less than three times length, greath ()\erhanging metanotiim (for one-half its length) (.Africa, southern /\sia) Paclnrinthidnini (part) 1. juxtantennal carinae present; T7 broadlv truncate with small median projection (Palearcdc, Africa, Oriental) — Eoarithidium (part) -Juxtantennal carinae absent; T7 with three apical spines (Africa) Snripista

COMMENTS, DESCRIPTIONS, .AND KEYS TO Acanthidium batrae new species SUBGENERA (Figs. 1,'_'. 7,8. 10-14)

Male: Bod\ length 1.3 to 8.0 mm; forewng length 6.0 mm: head The following pages consist of comments and descriptions width 2.8 mm. Hmd: Without carinae but preoccipital ridge be- hind vertex nearh flat in justifiing or explaining decisions made in preparing the List strongly angular. Clhpeus profile, upper margin between subantennal sutiues convex; lower lat- of Genus-Group Names. In addition, keys are given to the strongly eral short, rcflexed; lower concave, with that we have Genera are margin margin slightly subgenera recognized. arranged three small dark denticles, not overhanging base of labrum. to facilitate reference. alphabetically ready Mandible .^-toothed, interspaces between apices approximatelv equal, outer surface punctate, shining, with carinae evanescent in basal half. First .segment of labial palpus slightly shorter than sec- Acanthidium new genus ond: maxillan' palpus longer tlian greatest width of galea, proba- bly 4-segmcnted. second and third segments cylindrical, fourth (or Type species: Acanthidium balrae Michener and Griswold, new species. apex of third) minute and tapering. Subantennal sutures straight, longer than diameter of antennal socket, parallel, lower ends aris- This is a genus having the body form of Eoanthidium or ing from tentorial pits. Interantennal distance twice antennocu- Pseudoanthidium, and The conspicuous yellow markings. lar distance; ocelloccipital distance equal to interocellar distance, basal part of the axilla is produced lateralh and pointed, about nvo-thirds of ocellocular distance, (ienal area much narrower than eve seen from widest helou middle of not suggesting the shape of the axilla of some Rhodduthidium side, eye. Scape reaching level of anterior margin of anterior ocellus; first flagel- (Asianthidium) that led to the naming oi Axillanthidium Pas- lar segment nearly 1.,t times as long as broad, second slghtlv teels, and likewise its in Stelis (Malan- suggesting shape broader than long, third about as broad as long, subsequent seg- tliidium) malaccensis (Friese). Thus this striking character ments progressively a little longer so that tenth is conspicuously than broad, eleventh about 1 .d times as as broad. Tho- has e\identl\ arisen at least three times. If one ignores it, longer long rax: Without carinae except for strong carina on pronotal lobe; Acanthidium still does not fit into an)' other genus. The half of omalus but not carinate; foveate scutoscutellar suture Eoanthidium and upper sharply angulate quite suggests front end of scutum gradually cmyed down, without smooth ver- Bathanthidium. From the first it differs bv lack of jtixtantennal tical surface; axilla with margin near base produced laterally as carinae and the simple S6 of the female, among other fea- strong prominence behind which margin is concave; tegula widest medialh-; scutellum with broadly comex seen from tiues; from the latter it differs bv the lack of the post- posterior margin above, margin not caiinate, laterally (along with axilla) over- spiracular fovea of the propodetmi and the absence of the hanging, medialh' onlv slightly overhanging metanotum; scuto.s- row of across the base of the From all other pits propodeiun. cutellar suture forming narrow ftnea, wcakh dix'ided niedialK', fovea genera Amnthidinm differs in the short, broad T7 of the male easih' hidden b\' long hairs, pollen, etc. Propodeum without row of across base, laterally a doubtful indication of one or two with five apical projections, the lateral ones broadlv rounded, pits small fovea behind absent, fainth indicated the sublateral ones low, and the median one slender biu pits; spiracle possibly bv ridge close behind s|)iracle; profile of convex, bliuit. Carinae are on the head and thorax propodeum lacking except upper third declivous but not \ertical, cumng gradually to verti- for a carina on the the half of the oma- pronotal lobe; upper cal lower two-thirds. Front and mid basitarsi each aboiu as long lus is sharplv angular but not trid\' carinate. The sciuelhun as remaining tarsal .segments together, hind basitarsus shorter is than all basitarsi much shorter than tibiae; rounded posteriorK . not emarginate. roiuided in profile, remaining segments; hind basitarsus less than three limes as long as broad. Front and and scarcel)' overhanging the metanotimi. Sterna of the male mid tibiae each with one apical spine; hind tibia with apex lack oblique, combs. st convex medialh ; tibial spurs rough cuned at apices. .Arolia pre- This genus occius in India. The single known species is sent. Metasoma:T2 widest. Tl with line margining basal conca\'it\' .4. biitrac disunct only in middle third, horizontal surface more than half nnv spiries. as long as vertical siuface; Tl to Tb with posterior zones scarcely Etymolog)'; Akanthos, Greek, thorn, plus Anilitdiniii. with recognizable, feeblv depressed, somewhat more so on T4 and T5; reference to the basolateral projection of the axilla. tergal graduli ending near spiracles except perhaps on T(i which has lateral longitudinal carina; T6 othei^wise immodified. T7 306 The UNi\T.RSir\' of K\ns.\s Science Bueletin Oi.n World A\TMinrr.\'i-: Bees 307

several times as wide as lateral long, short, broad; margin produced As indicated by Bingham's (1898) description and ilhrs- as rounded lamella that forms median apicolateral lobe, projec- tration and a specimen in the Natural Histor\' Museum, Lon- tion a bknit, black-tipped spine slightK exceeding lateral lobes; don, Acanthidium /w/zr/c resembles siiperficiallv in form and between spine and lateral lobe is weaker rounded projection; me- coloration Anthidium desidiosum also from India. dian third of dorsal surface of T7 with strong transverse ridge. S2 Bingham, to S5 with posterior margins weakh and shallowh' conca\e, mar- A. halrae'is. however, smaller and entirely different in struc- zones smooth, S4 with ca- ginal impiinctate; strong longitudinal ture; for example, it has 4-toothed rather than 9-toothed rina siiblalerallv; S6 with hairv lateral shoidder and broadh mandibles in the female. rounded, translucent midapical sin"facc projecting somewhat be- yond shoulders. Puiiclattoii: On head and thorax dense, contigu- ous throughout including propodeinn, slightlv less dense on Genus Afranthidium Michener metasonia, less dense and finer on legs; tibiae not at all tubercu- late; apical depressed zones of Tl to T5 slightlv more fnielv piuic- This genus contains a large group of relati\es oi Anthid- tate than rest of terga. Piibesceyjce: Rather abimdanl and long on head and thorax and base, sides and venter of meta.soma, white, ium, generally rather small and robust, with multidentate slightlv dusk\ middorsallv on metasoma; terga except laterallv mandibles in the female as in Anthidium. Some species with hairs short, suberect; basitarsal hairs than di- longer longer agree with Anthidium dho in the depressed, medially widened ameters of basitarsi, but few such hairs on hind basitarsus. long zone of to for piemarginal T5 (see ke)' genera more details) ; Integinncnl: Black with \ellow markings as follows: mandible except the anterior margrin of this zone, however, is not angiilate for black apical margin and teeth, clvpeus, paraocular area ex- as in Anthidium. In other there is such tending as stripe almost to summit of eve; gena compleieh below species no zone, and and extending as stripe across vertex behind ocelli, pronotal lobe, the impiuictate marginal zone is sometimes broad, as in the anterolateral mark and lateral on .sciuimi, axilla, mediallv stripe siibgemis Immanthidium. These and the other external char- interrupted marginal stripe on sciUellimi, quadrate mark on acters indicated in the key to genera are generally distinc- mesepisternum below pronotal lobe, irregular area on mesoster- tive btit have For Anthidium nimi, legs except blacki.sh \entroapical areas on femora and most exceptions. example of ventral or inner sin faces of tibiae, broad transverse bands on (Proanthidium) oblongatum (Illiger) has T6 of the female Tl to T.T narrowlv interrupted mediallv with semicircular midlat- denticulate, without a lateral tooth, and with a small mid- eral posterior black intrusions into bands on Tl and T2; T(i. T7. apical notch, thtis combining the usual feattires of Anthid- and sterna whollvvellow. Posteiior margins of terga brownish, light ium and The wide of the brown on T5 and T6. Wings dusky, xeins black. Afrantliidium. separation penis valves of Anthidium the Ff,m\lk: Agrees with description of male except for usual sex- and long bridge between their bases ual characters and as follows. Head: Lower of margin clvpeus is the only known invariably good character that distin- for denticles which are as in male; mandible 4- straight except guishes Anthidium from Afranthidium. toothed, upper two interspaces gentl)- conca\e, outer surface finelv Afranthidium contains diverse elements and one cotild pimctate, especialh apicalh , carinae distinct onlv on apical third. (Proboscis not examined.) Interantennal distance less than twice justih' di\iding it into several genera. Lhitil a proper analy- antennocular distance. First flagellar segment less than 1.5 times sis is made, we have not done so. The subgenera Imman- as as broad, second almost tivice as broad as long long, following thidium, Nigianthidium, and Zosteranthidium are particularly but ninth still broader than segments progressively longer slighlh disUnctive. long and tenth about 1 .5 times as broad as long. Tlinnix: Front ba- sitarsus slightly longer than remaining tarsal segments together, other basitarsi about as as Mflasi>iiia:Tli long remaining segments. Key to the Subgenera of Afranthidium to T4 of about equal width. T6 without lateral carina, slightlv con- cave in profile because of elevated apical area, margin with small 1. Margins of T2 to T:5 broadlv impunctate. median midapical notch; S6 unmodified, slightK' exceeding T6. Pubes- of zones tisuallv one-sixth cence: On dorsum of head and thorax with intermixed dusk\ hairs; lengdis impunctate length of sides of metasoma without long hairs; scopa yellowish white; ba- exposed parts of terga or more, margins trans- sitarsi with few hairs longer than diameters oi basitarsi excejjl parent, pale brown or cream colored; male gonos- lower of hind basitarstis with manv stich hairs. bUcgiimnil: margin t)lus enonnoiis, broad, flat, and almost membranous; Frons below median ocellus with vellow mark; and upper poste- lobe not carinate, scjmetimes with small rior stuTaces of femora with black areas; \ellow bands of metasoma pronotal and hain in of carina without midlateral black intrusions; S6 largelv black. ptinctate ridge position Holot\pe male: INDIA: Uttar Pradesh: Mussoorie I.al Tibba, al- hnmantliiilium titude 7.500 feet 23. 1963 W. T. on "In- (2:^08 m),June (S. Batra), —Margins of T2 to T.5 pimctate or narrowlv impunc- digofera dusua Buch.-Ham." Paratvpe male: Same data but Juh 1, tate, median lengths of impiuictate zones about 1965. Paratvpe female: Same data but June 25, 1965. These spec- one-seventh of of or less, imens are in the Snow Entomological Museum, University of lengths exposed parts terga dark or translucent Kansas, Lawrence, thanks to the generositv of Dr. S. W. T. Batra, margins brownish; male gonos- after is whom the species named. tyius not broad, flat and almost membranous al-

Figs. 1-6. Genitalia of males, dorsal-ventral and lateral views; in the former, dorsal views are at the left, ventral at the right. Let- ters identifi.' same structures for any one species, e.g., a in a dorsal \'iew is the same as a in a lateral view. I, 2. Acanthidium balrae Michener and Griswold; sclerotization that may be in the basal part of die endophalltis is shown at left in dorsal view, and at lower right of Fig. 2, in lateral view. 3, 4. Indautliidium crriudalicauda Michener and Griswold. 5, 6. Pseudoaiithidiuni (Tul)erantliidium) brachealum Michener and Griswold. Figs. 7-9. Apices of mandibles. 7. .4(y/»//»V//(/h/ /w////(c Micliener and Griswold. male. 8. S.ime, female. 9. Indantliidiuin civiiuldti- cauda Michener and Griswold, male. 308 The Unatrsityof Kansas Science Bulletin

Figs. 10-14. Acanlhidmm t/rilim'Mii.\\cnvr dndGrhvio\d. 10. Face, female. 11. Face, male. 12. Slightly lat- eral view of dorsum of thorax, male, to show shape of axilla (a). 13. Metasomal apex, male, slightly lateral view to show two convexities lateral to median spine of T7. 14. Forewing. male.

though sometimes bioadiv paddle-shaped; prono- hanging metanottim; lateially scutellum and usually tal lobe usually with transverse carina or km lamella a.xilla overhanging large fcjssa and iisualh aciUe or which is hairless even when it extends only partwav narrowly rounded as seen obliquely to show pro- across lobe file of lateral part of scutellum and of axilla; outer Scutellum rotnided in profile, not or scarcely surfaces of tibiae, especially of female, coarsely, ir- oxerhanging metanotum, even laterally axilla and regularly pimctate, sometimes with extensive scutellum not much overhanging; outer surfaces of smooth areas between pimctures, these ptinctures tibiae not more coarsely punctate than mesepister- commonly coarser than those of mesepisternuiu, num, not tuberculate tibial surfaces usually tuberculate, not or weakly so -Scutellum acute, right angular or sometimes in Cajmulhidium and Mesanthidium in rounded profile, medially often strongly over- 3. T2 to T5 with apical bands, broken medially, of Oi.n \A'()Ri,n A.v77//D//.\7; Bees 309 — white plumose hair; propodeal triangle with punc- Posterior margins of at least some metasomal terga, tures (and hairs) widely separated from one an- seen in profile, cm"\ed upward away from following other, surface between punctures strongly terga; axilla not extending laterally beyond sciumn shagreened, dull; boch and legs black, withoiu pale except in some Bmuthidiitm; male gonostsius reach- markings Zosteranlhidium ing to or beyond level of apical fourth of length of —T2 to T5 \vitlu)iu apical hair bands [except for penis valve 8 Afrauthidiuin murinum (Pasteels)]; propodeal tri- 8. T6 of female with preapical denticulate ridge angle rather denseh punctate and haiiT, surface parallel to denticulate apical margin (sciuelhun between pimctures shining; bod\ and legs usually distinctly carinate except for small midapical emar- widi yellow or white markings, but if not, then at least gination) (male unknown) Xenanthidium anterior surfaces of front and mid tibiae brownish —T6 of female without preapical denticulate ridge ... .9 yellow 4 9. T6 of male with preapical usually denticulate trans- 4. Hind basitarsus of female with apical projection verse ridge at least laterally; tibiae coarsely punctate oyer base of second tarsal segment; hind trochanter but not or weakU tuberculate on outer surfaces of male with apicoyentral denticle; sterna of male Capanlhidium with distinct basal fasciae arising at graduli; body —T6 of male without preapical ridge; tibiae strongly black, \ellow or cream color usually limited to tuberculate on outer siufaces 10

minute streak along inner margin of eye of both 1 0. Preoccipital carina present laterally; male S3 with sexes Nigranthidium trapezoidal apical projection; T5 and T6 of male wAxh —Hind basitarsus of female Mth apex truncate, with- lobate lateral carinae; female T5 and T6 with lateral out apical projection; hind trochanter of male with- longitudinal carinae Mesanthididlum oiu apicoyentral denticle; sterna of male without —Preoccipital carina absent; male S3 margin not pro- basal with or cream colored lateral fasciae; body yellow duced; T5 (ustiall) ) and T6 without carinae areas at least on face of male and metasomal in either sex Branlhidium terga' 5 5. T5 of female without lateral spine; Tl of female Michener s. str. with carina separating anterior from dorsal siufaces Subgenus jyranthidiiim abrupd)' sd engthened laterally and thence extended The species described as Anlhidioma muniiiim Pasteels lateroposteriorly; apex ofT7 of male with two lobes, (1984) is not closely related to Anlhidioma and appears to each two or three times as broad as long, emar- belong to the genus Afranthidiuvi, but does not completely gination between them with small median spine; S6 fit the characterization of any recognized subgenus. As the of male with small, pointed midapical process at base species is known onl\- in the female, a firm decision as to its of which are two spines directed forward is The lobe lacks a carina Domunlhidium placement premature. pronotal but in other respects murinum runs to couplet 3 in the key —To of female with lateral spine; Tl carina of female to genera, or to Afranthidium s. str. if its tergal hair bands luimodified; apex of T7 of male 2-lobed, usually with are ignored. It differs from Afranthidium &. str. in the absence median angle or tooth, or trifid, lateral lobes about of lateral metasomal spines although T3 to T5 ha\e small as long as broad or longer; S6 of male withoiU small lateral lobes, and from Zosleranthidium in the presence of apical process and spines (i \ ellow maculations and other characters. Afranthidium mur- 6. Gonost\lus of male tapering, not or scarcely inum (Pasteels) (new combination) is unusual in its abun- longer than gonocoxite, not attaining middle of dant white hair, which forms dense apical tergal bands on penis valve; ventral surface of mesepisternum of fe- the metasoma. suggesting Zosteranlhidium. The species is male covered with sDong, backward directed, golden known only fiom Namibia, hi addidon to the t\pe specimen, to black bristles that appear flattened, miniuely two additional females from approximately the t)'pe local- barbed, and bkuit or abrupth tapered at apices (S6 it\' 1 (Pomona, in Diamond Aiea No. , Namibia) were taken of male elongate, produced to narrowl)' roimded or on flowers of a yellow legiune b\' V. B. Wliitehead and are bidentate apex) Oranlhidiitm — in the South /\frican Museiun, Gapetown. Gonost\lus of male expanded apicallv. about twice as long as gonocoxite, attaining apex of penis valve or nearly so; ventral siuface of mesepistermmi of fe- Subgenus Branthidhim Pasteels male with pale, gradually tapering hairs similar to those of adjacent areas Afranthidhnn s. str. Honanthidium Pasteels (1969a) was described as near Tu- 7. Posterior margins of metasomal teiga not ciu^ved heranthidium (here considered a subgenus of Pseudoanthid- upward, hi]ig near surfaces of following terga; ax- ium) and was later SMiouMnized with Tuberanthidium (Pasteels,

illa extending lateralh' beyond scutal margin; male 1984) . It is known onh' in the female, so its place is not read- gonost)ius greatly reduced, attaining aboiu middle ily determined with certainty. However, it seems to us much of penis valves, which are completely fused to one more likeh to be an Afranthidium, closest to the subgenus another (palearctic) Mi'sdiithidiinii Branlhidium. to which it runs in the key to subgenera. 310 The University of Kansas Science Bulletin

because of the strongly denticulate (and not emarginate) Subgenus Xenanthidiinn Pasteels T6 of the female, the presence of weak swellings at the Xenanthidium should be considered a sides of T2 to T5 (these could represent spines of the male; probably synonym of but since it is known from a fe- they are not present in females of Pseudoanthidkun) and the Capanthidium, only single rather narrow, unbroken vellow bands of the metasomal male specimen that has rather distinctive characters, it seems to it. terga. Unusual features are the swollen head, with the genal premature synonymize A folded label on the oi X. Insnra- area broader than the eye, and the strong and elevated large type specimen tum Pasteels trimmal carina extending from the mandibular acetabulum (Natural History Museum, London) provides much better data than did Pasteels in to the fourth mandibular tooth. This is probably not ho- (1984) print. Com- this folded label with the label that Pasteels the mologous to the swelling in the same region, but without bining read, a carina, found in Tubmnilhidium. The fifth (small) and sixth type is from Pouss on the Logone River in north Cameroon, 200 miles south of Lake Chad, 250 Novem- (uppermost) mandibular teeth are depressed, so that Afmn- m [altitude?], tliidium (Branthidium) hoj^estum (Cockerell) (new combina- ber, 1979 (G.Popov). tion) runs with difficulty to the multidentate part of the key to genera imless the mandibles are opened. Teeth two to Zosteratitliidium new four, however, are of more or less equal size, separated by subgenus narrow notches as shown Pasteels some- (Figs. 1.5-17) by (1969a, fig. 82) ;

how a tooth was lost in Pasteels" (1984, fig. 124) subsequently Type species: Nigifinllu/liinn Ingofcnnnlinti Pasteels. 1984. later drawing. Afrdiithidium (Branthidium) i^iillaniKirli (Maxromoustakis) Although the type species of Zosteianthidium was placed is in Pasteels it is not related as identified by Pasteels unusual in having the axilla ex- Nigranthidium by (1984) , closely to that Its tending laterally as in the subgenus Mesanthidium and T5 subgenus. somewhat elongate body, pale tergal (also T3 and T4 but not T6) with a lateral, almost carinate hair bands suggesting Afranthidium nuuinum Pasteels (see lobe in the female, suggesting MesanthidicUum which has such discussion oi Afranthidium s. str.), and complete absence of carinae on T5 and T6. yellow or white integumental markings, as well as its size (body length 9 to 10 nun), result in a species that exactly resembles some species oi Hoplilis (Osmiini). Its distinctive Subgenus Capanthidiutn Pasteels features are as follows: MaxillaiT palpu.s as long as widlli of maxilla at point of palpal Anthidiuw ior Type species: "lYifjHdh'Fvwse," lapsus lapuoht Biamis, 190.5. attachment, second segment about seven times as long as wide, Pasteels ( 1 969a) twice lendei ed the name by original designation. specific widest near ba.se, tapering, bristly (thus maxillaiT palpus longer once and each lime attiibnied ii to Fi iese, in whose lapiiole, tapkola, than in any olhcr Afrtiullndiiim): labial palpus with second .segment Brauns' was paper species published. about twice as long as first, third segment broadly attached to sec- ond and continued in same direction, onl\ loitrth directed lat- This includes certain at least subgenus palearctic species, erally. Mandible of female 6-loothed, teeth 2 and 6 sitbequal, 5 A. (C.) noeft (Benoist) and srhulthessii (Friese), formerlv smallest, teeth 4 and .5 a little shorter than the others but apices of all teeth the same line; tnandible of male 3-toothed. placed in the subgenus Mesanthidium. The male genitalia nearing Clypeal apex denticulate in female. Second recurrent vein slightly of the seven or more additional palearctic species placed beyond second transverse cubital; stigma with distal half taper- in Mesanthidium Pasteels by (1969a) presiunably pro\ide ing into marginal cell. Sciitelluni roitndcd posteriorly, not at all the most reliable basis for these in Mesan- tri- placing species overhanging; ]3i tipodeum strongh shagreened, punctures on almost absent in median zone and lower of thidium (with verv reduced gonostyli, see key to subgenera) angle dispersed, part triangle, hairs con espondingly sparse. Hind basitarsus with apex or Capanthidium. truncate, not produced as in Nigraulliidium. T2 to Tb with preapi- cal bands of xvhite plumose hairs, broken middorsalK'; T5 with pos- terior premarginal zone depressed but not sharply defined, Mesanthidiellum Pasteels Subgenus somewhat more finely punctate than test of teigum, anterior margin of zone not angulate medially (so poorly defined that angle One of us (CDM) would consider this a synonym oi Btan- might not be visible), narrowed laterally but more than half of median width, black, broader than tliidiiim. impunctate tergal margin base of last hind tarsal segment; T(i of female without preapical carina, profile straight, reflexed lateial portion smaller than in related bees, exposed reflexed area being about as long as dis- Pasteels Subgenus Nigt-anthidium tance from distal end of area to small midapical teigal notch, thus about half as long as in other A/raul/iidinm. Tfi of inale with lat- eral tooth; T7 of male not exserted, with broad emar- Pasteels (1984) included two species, A. (N.) roneolor strongly between two teeth as illustrated Pasteels (1984). (Friese), of which Osmia unllowmorensis Brauns was consid- gination by Metasomal sterna more modified than in other Afranthidium; S2 ered a and A. (Pasteels). The synonym, tergofasciatum syn- and S3 with preapical zones of veiy long white hairs, S3 with mid- of O. willoiummensis is incorrect. It thus forms the onyTny new apical area of stiff, straight hairs; S4 with large midapical area of Stiff, S3 with small lateral tubercle combination A. (N.) willowmorense (Brauns). A. lergojasciatum straight, brown hairs; concave, or tooth, and hairless (Pasteels) is transferred to the subgenus Zosteianthidium. posterior part shining except posterior marginal band of short straight brown hairs on lateral third of Old World Axrnmii\E Bees 311

15. male. 16. female. Figs. 15-17. Afraiilhidium (Zosleianlludium) Ingofasciatum (P:\steeh). Mandible, Mandible, 17. Portion of forewing, female. male. 19. female. 20. Figs. 18-21. Iiidaulh/Hiiim ireniilfilirauddMichener RndGrhwo\d. 18. Face, Face, Forewing, to male. 21. Metasomal apex, male, slightly lateral view to show teeth of T7 lateral median spine (m).

sternum: S6 broad, shining, largely impimctate and hairless, with tains rather diverse forms. In the difficult couplet 2 of the lateral thin, broadly conyex; strong gibbosity, posterior margin key to genera, Zoslcmnthidium runs to 4 because the pre- S7 broad with apical projection margined by hairs [this shape is marginal zone ofT.5 is poorly defined, not greatly narrowed 1 1 as . slender, illustrated by Pasteels ( 1984. fig. 4) S6] Gonostylus its anterior not and the .smooth about shape shown b\' Pasteels (1984) biu with seyeral large laterally, margin angulate, broad. cuned hairs arising on upper and inner surfaces; penis valve posterior margin relatively with apex bent upward and subtruncate. Zostrranthidium contains a single species, Afranlhidium is distinctive that it could well receive This taxon so (Zostennilhidliim) tergofasciatum (Pasteels), new combina- generic status. It does appear to be nearest to Afmtithidiuw, tion, from western Cape Province, South .Africa. and for the is included. As indicated elsewhere, the present EtvTOology: Z.oster, Greek, belt, with leference to the meta- is for it con- status of Afranthidvmn considered tentative, somal hair bands, plus Anthidium. 312 The UxiMRSi n of K\nsas Science Bulletin — Genus Afrostelis CockereU Tegula broadly roimded or almost transverse pos- teriorly; omalar carina extending to venter, or in The two on front and middle tibiae and other spines some Pyenanthidium very weak or perhaps absent on characters indicate that is related this genus closely to Stelis; lower half of mesepisternum 5 it be a cleri\ilive of Stelis not war- might merely specialized 5. Tl with carina separating anterior and dorsal rank. However, the male form ranting generic gonostvlar surfaces; apex of mandible but little wider than (slender, and minutelv is more straight, capitate) probably base; body black with yellow or cream markings than that of Stelis. That fact and the primitive any striking on all tagmata (Africa, soiuhern Orient, northern thoracic characters lead us to as a recognize Afrostelis genus. .Australia) Pyenanthidium —Tl withoiu carina between anterior and dorsal sur- of mandible of female about 1.5 times Genus Anthidiellum CockereU faces; apex as wide as base; body brown with red markings or yellowish and black but without on Tl to T4 Key to the Subgenera oi Anthidiellum yellow (southeast ,A>ia, Indonesia) Ra)ithidiellum

1 . Postgradular parts of T2 to T5 of females and to T6 of males swollen so that from above laterally Subgenus Clypanthidium Pasteels sides of metasoma seem lobed; S.T of male without DiscoveiT of the male of this comb; mandible of female with preapical shoulder subu;enus misrht show that it is oiU of in on lower margin, below lower tooth (Holarctic) place Anthidiellum. The Ajilhidiellum-Mkt; fea- tures it Anthidiellum s. str. of Clypanthidium, by which differs from Bathanthid- — /((w where it was Pasteels include Terga not swollen laterally; S5 of male with margin placed by (1968, 1969a),

( 1 the of a carina on the half or of broadly concave and armed with comb of black ) presence upper more the omalus extends farther down in other teeth, at least laterally (male imknown in Ananthi- (it Anthidiellum), the diellum); mandible of female without preapical (2) enlarged sciuellum, strongly overhanging the meta- notimi it is not shoulder on lower margin (eastern hemisphere) (but sharply angled, carinate or lamellate api- 2 cally, as in other Anthidiellum), (3) the rather abrupt line between the area with keirotrichia and that with other 2. Mandible of female miniuely sculptured, dull, al- hairs most without carinae, apex expanded, 1.5 times as along the upper margin of the hind tibia (biU it is not so as in otlier die robust hind basitarsus wide as basal width; subantennal siuure straight; abiTipt Anthidiellum) , (4) is about three times as as about as in Tl to T4 without pale markings (southeast Asia) (it long wide, Anthi- Ananthidielluin diellum. but is over four times as long as wide in Bathanthid- — and the robust form. Mandible of female somewhat shining, often coarsely ium) (5) body pimctate, somewhat dull distally in Ranthidiellum and Clypanthidium, with carinae, apex but little wider Subgenus Pyaianthidium Kronibein than base except in Ranthidiellum; subantennal su- ture arcuate [scarcely so in A. (Anthidiellum) bre- The name Pygnanlhidiellum was proposed for the .African viusculum not (Perez); clearly recognizable and species and contrasted to the Indoaustralian species. In the not arcuate in A. bimacula- perhaps [Clypanthidium latter group the hind tibia and basitarsus are finely punc- tum (Friese)]; Tl to T4 with vellow, cream, or red- tate, the omalar carina is weak or absent below, and the dish bands except in subgenus Cl\panlhidinm and pronotal lobe is carinate or weakly lamellate. In the .African Ranthidiellum some 3 group the hind tibia and basitarsus are coarsely punctate, 3. Preoccipital carina present at sides, behind eyes, the omalar carina is complete and the pronotal lobe some- but absent behind vertex; lamella on pronotal lobe dmes strongly lamellate. Also in the African group the hind tapering but extending nearly as far mesad from lat- basitarsus of the female is enlarged, nearly as wide as the eral margin of sciuum as laterad; axilla extending tibia. It now appears that both groups occm^ in Sri Lanka,

laterally beyond margin of scutum (basal zone of India and Binma (Pasteels, 1972) . We do not regard the dif- horizontal with propodeum laterall)' well-developed ferences between these groups asjustification for subgeneric —pits) (east Africa) Chloranthidiellum distinction. Preoccipital carina complete, present only behind vertex or absent; lobe with lamella or ca- pronotal Subgenus Ranthidiellum Pasteels rina usually extending little mesad from lateral mar- of Pasteels gin scutum; axilla not extending laterally beyond designated the type species as "Anthidium rufo- lateral margin of scutum 4 maculatum" Cameron, and on p. 123 he described it, in- 4. Tegula narrowly roimded posteriorly; omalar ca- cluding the reference "Cameron, 1897, Mem. Mancliester rina reaching about half\vay down omalus (south- Soc, 41." Cameron published in 1897 in \'ol. 41 of that east Asia) Clypanthidium journal, but Anthidium ntfomaeululum does not appear there. Old Wt)Ri.D AMHfDfixi-: Bees 313

It is not the as same AnthuUum ru/oiiuirnlaliiiii Friese, 1899, tate medially, but laterally it is as described. In A. (Severan- from There is, FioliKinlhitliui/i Syria. ho\ve\er, riilniiianilritiiiii thidium) seven niy-cichA, also from Africa, the punctures of Cameron, 1902 (Jour. Straits Branch, Ro)al <\siatic Societ), the depressed zone are sparse and shallow and this zone no. that 37) agrees reasonably well with Pasteels' descrip- merges into the smooth margin; the shape of the depressed tion. No doubt this is the that Pasteels intended is species as zone, however, as in other Anthidium. The preapical de- the oi type species RanthidicUiim. pressed zones are wider (at least on T5) medially than lat- alternati\e at An way of lookins^ the t^pe species of Raiilhi- erally; this is usualh e\ident e\en when the zone is not well diellum is to list it as Anthidiellum mjumaculutum Pasteels, 1969, differentiated medially. The anterior margin of the preapi- by original designation, recognizing that his attribution of cal zone is usually angled medially so that the whole zone the name to Cameron the were errors. is a (and reference) very broad triangle. These characters of the tergal mar- Pasteels described the species in full, as though it were new. gins are best examined on T5 of both sexes, but are often Anthidiellum latum seems to be a nifomani Pasteels, 1969, ewdent on more anterior terga. In the other genera with of, as well as Pro- similar synonym ajimior secondary homonym of, female mandibles, the smooth apical tergal mar- toanthidium nifomaculatumCan\eron, 1902. Thus ultimately gins are usually convex and the preapical zones are not rec- the of RarithidicUum is the same as is type species indicated ognizable except laterally or are scarcely wider mediallv

in the . In preceding paragraph. than laterall) some species of the subgenus Proanthid- ium the marginal zones are also convex. Genus Anthidioma Pasteels

Key to the Old World Subgenera of Anthidium In the absence of males, the true position of Anthidioma is not clear; for the its rank is maintained. present generic 1. Scutellum roimded in profile, not carinate or Pasteels (1984) included two each then known species, lamellate, not greatly overhanging metanotmn and from a female in Anthidinma. One of them, single specimen, propodeum; pronotal lobe with or without carina A. murina Pasteels, is an see the discussion Ajranthulium; (Holarctic, Oriental, Africa) Anthidium. s. str. under s. str. One additional female of an An- — Afranthidium Scutelhun angulate in profile (at least as seen thidioma related to but different from A. chali- species obliquely to show profile of lateral part of scutellum), codomoides Pasteels has come to hand. carinate or at strongly lamellate least laterally, greatly overhanging metanotum and propodeum; pronotal Genus Anthidium Fabricius lobe carinate or lamellate 2 2. Pronotal lobe carinate; axillar suture weak; scu- The distinctions bervveen Anthidium and the remaining tosciUellar siUure not in deep depression so that genera (as a group) with tapering mandibular teeth in the scutellum nearly continues profile of scutum (scutel- female are rather subde and tend to break down among prob- lum strongly produced posteriorly as rather flat ably derived subgenera that seem to ha\'e lost one or another structme ending in lamella) (east .Africa) of the characters of the genus. The combination of char- — Nivanthidium acters remains distinctive, however. In Anthidium the sub- Pronotal lobe with more or less anteriorly directed antennal suture is usually straight. T6 of the female has an or erect, translucent lamella; axillar suture strong; apical depressed rim, usually smooth and shining, often hid- scutoscutellar suture in depression so that scutellum den by hairs, and sometimes imrecognizable. This rim and is independently convex in profile 3 usually the tergmn as a whole have a median apical notch 3. Scutellum ending in lamella or large carina al- or all emargination, sometimes small or largely hidden by most the way across; hind basitarsus with longi- hairs, but sometimes large and conspicuous, especially in tudinal carina on outer surface; omalus sharply the Callanthidium. T6 of the or carinate 4 subgenus Laterally, female —angulate weakly nearly always has a tooth, angle, or shoulder, mesal to which Scutellar margin with broad median part neither there is an emargination, sometimes veiy weak. In the sub- carinate nor lamellate: hind basitarsus not cari- genus Proanthidium the lateral emargination and tooth are nate; omalus roiuided or formina; rounded angle absent but the impressed margin and notch are present me- (Palearctic) 5 dially. T1-T5 of females and T1-T6 of males have narrow, 4. Posterior scutellar margin seen from above sub- smooth apical margins of uniform width, usualh' flat or truncate, its lateral part ciuAed fonvard becoming nearly so. Anterior to each margin but behind the elevated more or less longitudinal; antennae below level of middles mid-tergal zone (often distinct only laterally) is the de- of eyes which converge suongly below so that pre.ssed marginal zone, differentiated (commonly by finer clypeus is imusually small (Africa, Arabian Penin- and closer punctaUon) from the rest of the tergmn. Prob- sula) Severanthidium lems this — with character are foinid, among others, in A. (Nii'- Posterior sciUellar margin seen from above broadlv uuthidium) uiveocinctum Gerstaecker from Mrica, which roimded with small median emargination, laterallv has the depressed zone of T5 of the female sparsely punc- oblique, only at extreme end next to axilla some- 314 The University of Kansas Science Bulletin

row of times becoming longitudinal; antennae near level without pits except laterally) (southern (Jhina, to mod- Bathanthidium s. str. of middles of eves which converge slightly —Taiwan) erately (xeric southern Palearctic) Gulanthidium Fovea behind propodeal spiracle elongate, weakly 5. Scutellum seen from above with margin curved for- delimited; T7 of male simple or with slightly pro- ward at each side to axillar margin; hind basitarsus duced median lobe 2 of female less than four times as long as broad (cen- 2. Basal zone of propodeum not distinct, with ir- T(i tral Asia) Turkanthidiinn regular, sculptured, sloping pits lateralh'; of male —Scutellum seen from above with margin more or less with elevated median section and concave lateral sec- transverse, curved foi-ward rather abruptly or angled tions (Oriental) Manlhidium foi"ward at side to axillar margin, scutellar margin —Basal zone of propodetun distinct laterally with reg- horizontal of male with often with lateral angle protruding posteriorh : hind ular, shiny, nearlv pits: T6 basitarstis of female foin- or more times as long as surface convex (northeastern Asia) . . .Stenanthidiellum broad (Palearctic) Proanthidium

Subgenus Stenanthidiellum Pasteels Subgenus Anthidiiim Fabricius s. str. Romankova (1988) considers the type species oi Stenan- diverse Unusual This is a large and rather subgenus. thididlum and Lasanthidium (listed above as a probable ju- or species or small groups have been given subgeneric nior synonym of Stenauthidirllum) to be in different genera. but such names seem generic names, unnecessary, being ap- The identity oi Anthidium .\ihiri(um Eversmann (t)'pe species on derived from the "ordi- parently based species among oi Stenanthidiellum) may be uncertain. The genitalia oiStelis oi Anlhidhun s. str. nary" species malaisei Fopov (type species of Lasanthidium) are similar to Ardenlhidium has been to a of The name applied group those oi Bathanthidium (Bathanthidium) hifoi>eolatum Miken about ten mandibtilar teeth in unrelated species having (see Popo\, 1941) and it may be that Batltanthidiitm and the female, compared to five to seven in most other species. Stenanthidiellum should be imited. In other respects the type species, A. ardetis Smith, and the dissimilar A. Friese appear to be An- fjuite undulatifonne Genus Benanthis Pasteels Ihidium s. str. A. echinatum Klug, the type species of Echi- uaiilhidiinn, is better differentiated from most other perhaps This genus, known from a Malagasy specimen that has ap- Aniliidiiini s. str. It is one of the few forms wilh- palearctic parently been misplaced or lost, is unknown to us. Pasteels oiU a hind tibial carina. The clypeal margin of the female and sketched of the ( I9(i9a, 1984) described parts specimen. is not thickened as in most Anlhidium s. str. In this respect We cannot comment on its probable relationships. it resembles Tarkanlhidium and most Proanthidium. Like to the manv desert bees ( Morocco Pakistan ) , species placed Genus Pasteels in Echitianthidium have a pallid aspect clue to the largely yel- Cyphanthidiuin low metasoma with preapical tergal fringes of dense white Two species were placed in separate genera (Cyphanthid- hairs. A. pontisCocVcveW (Puiiliuithidiiim) differs from other iumand Trianthidiellum) by Pasteels (1969a); Pasteels (1984) Anlhidium s. str. in the protuijeranl clypeus, a character transfeiTed Trianthidiellum to Anlhidiellum as a subgenus, per- that alone seems not to justify subgeneric recognition. haps because of the arcuate subantennal siUures. However, in specimens of "Trianthidiellum" near C. sheppardi (Mavro- Genus Bathanthidium Mavromoustakis moustakis) (type species of Trianthidiellum) (National Col- lection of Insects, Pretoria, South Africa) they are straight, This contains three taxa that have genus-grotip gentis and in C. sheppardi they are only slightly arcuate; Pasteels' names. Because only four species are involved, it may seem the siUiue ctuTed on one side figure (1984, fig. 240) shows imreasonable to We have done so recognize subgenera. but nearh' straight on the other. Ma\Tomoiistakis (1937) de- because there is ncj assurance that the is mono- parth' genus scribed C. sheppardi as lacking arolia but they are present The Bathanthidium s. str. and Stenan- plnletic. subgenera although rather small in the type specimen as well as in other thidiellum are slender bodied, close relatives: quite clearly specimens that we have seen. Thus there seems to be no jus- both have a small median comb on S4 of the male. The e.g., tification for tw(j genus-group names for the tvvo descriiaed subgenus Manlhidium is more robust, resembling Anthi- species. dicllum subgenus Ranthididlum. All three siibsrenera have a comb on S5 of the male almost the entire width occupying Genus Eoanthidium Popov of the .segment. Key to the Subgenera oi Eoanthidium Key to the Subgenera of Bathanthidium 1 . Subantennal suture straight; inner surface of hind tibia onto withoiU line 1. Fovea behind propodeal spiracle rounded, de- cuning upper margin sharp between keirotrichiate area and of limited by strong carina; T7 of male trilobed, me- region longer hairs sciUoscutellar siUiue closed, dian lobe longest (upper margin of propodeum (arolia present; Old World A-wrnrDiixi-: Bees 315

similar to scutoaxillar suture; T4 to T6 of female and spiracle, posterior margin clearly defined but lower end weak or and end or closed a T5 and T6 of male with small lateral spines) (south- open upper open by carina. Other features of in with ern India) HiiiiuUellum Clislanthidium, mostly agreement — some or all other subgenera of Eoaiilliidium, include the follow- Siibantennal suture strongly arcuate outward; inner ing: preoccipital ridge noncarinate; subantennal suture strongly surface of hind tibia flat, keiotrichiate area arcuate, lower end from tentorial ending arising pit; hind tibia with sharp line and carina abruptly at sharp line, often carinate, along upper separating iiuier. keirotrichiate surface from rounded suiface; without row of margin of tibia 2 upper propodeum pits across the upper margin except as suggested at the exueme sides; lateral meta- 2. Aiolia absent; profile of T6 of female convex; T4 somal teeth absent except lor tooth at side of T(i and S5 of male, to T6 with lateral sumre spines (scutoscutellar closed, small teeth also at sides of S3 and S4 of male /:. (C.) nasirum:Tl similar to scutoaxillar suture) Salemanthidium of male broad; sternal combs of male absent; T7 of male with me- dian but not carinate —Arolia present; profile of T6 of female concave point reaching extending beyond posterior T6 of female with or distallv in tergal margin; obtuse acute lateral angle; be- (convex subgenus Clistaiit/iidiiiin) ; T4 and tween sti these angles ong, denticulate premarginal ridge without T5 without lateral spines 3 median emargination; tergal margin proper inconspicuous, be- 3. Front coxa with carina or lamella trans- strong hind the denticulate ridge, simple except for small median emar- to axis verse of body or extended distally and thus gination. Clistanthidium is largely longitudinal; S6 of female not thickened api- known from Namibia to Natal Province, calh, withoiu lateral carina or spine; T6 of male South Africa, north to Shaba Province in Zaire and to Tan- with lateral tooth; scutoscutellar suture not veiy dif- zania, Mali, Ethiopia, Israel, Iran, Pakistan and Turkey. We have ferent from scutoaxillar suture, or in E. (C.) nasicum studied Eoanthidium (Clistanthidium) amiaticeps (Friese), (Friese) foveate nearly as broadlv as in E. (Eoanlhid- bituberculatum (Pasteels). rothschildi (Vachal), and tumer- iiim) s. str Clislanlludium icum (MavromoiLstakis) from Africa and E. (C.) nasicum —Front coxa without strong carina or lamella; S6 of (Friese) from Asia. female thickened apically, with lateral carina or Etymolog)': The Latinized form of the Greek klcistus, spine; T6 of male without lateral tooth; scutoscutellar closed, with reference to the closed scutoscutellar stiture, suture open with shinv impunctate floor divided plus Anthidium. mediall}' to form two foxeae Eoanthisium s. str. Genus Euaspis Gerstaecker Clistanthidhim new subgenus Euaspisis a paleotropical gentis of parasitic bees present in both Africa and Asia. The scutellum of oriental Type species: Diinilhiiliinn lunimium Mavromoustakis, 19;M. species differs from that of African species in form, being thickened, The species of this subgenus were placed in Eoanthidium punctate and bain in Oriental species rather than thin, lamel- {zs a subgenus oi Anthidiellum) by Pasteels (1984). Clistan- late and nearly hairless. On this basis, the Oriental species have thidium differs from Eoanthidium s. str., however, in several been recognized as a separate subgenus, Parevaspis Rit- characters that suggest a sepaiate subgeneric status. It is more sema (Pasteels, 1980). In the absence of other characters, such a robust, ha\ing the body form of a. Dianthidium. It differs from separation seems unnecessaiT. Parmaspisis therefore s. considered Eoanthidium str. in the presence of two blunt spines or a to be a junior synonym of Euaspis. single broad bilobed spine at the apex of the middle tibia, this condition is although approached in E. (Eoanthidium) Genus Gnafhanthidiiim Pasteels clypeare (Morawit/), and in the simple margin of S(i of the female, not at all thickened, although S(3 has a median keel Gnathiniliiidium is related to Pseudoanthidiitm and should ending in a preapical, median spiae. Clislanthidium seems perhaps be regarded as a subgenus of that genus, closest to the to be most similar to Eoanthidium s. str. from which it dif- subgenus Micranthidium as indicated by the strongly fers as follows. carinate, almost lamellate preoccipital ridge, pronotal lobe, Juxtantennal carina well developed, ocelloccipilal distance less omalus, and scutellar trimcation; the lack of lateral teeth than oceilocuku and interocellar distances, onialai carina (which on T6 and T7 of the male; and the bilobed T7 of the male. extends onto thoracic venter but does not approacli midline) el- It differs, however, in the nearly straight subantennal suture evated to form low lamella on upper half of niesepisternum; ca- and the not concave S3 to S5 of the with rina of pronolal lobe elevated to form low lamella that extends particularly male, no imtisual around lower end oflobe and upon posterior side; front coxa with setae except median (not marginal) patches of lamella transverse to axis of this lamella strong body, produced dense white hairs on S3 and S4, the posterior margins of to median in most males E. tiasirum anteriorly angle [in (C.) which are both convex, translucent, and hairless. (Friese) lateral part of lamella reduced and mesal part extending distad towaid coxal spine (male), so that lamella is laigelv longi- tudinal]; of axilla and scutellum for sh()rt median margin (except Indanthidiiim new genus section) produced as strong carina or lamella overhanging nieta- notum and propodeum: scutoscutellar suture nearly closed, simi- Type species: Indinilhiihinii ( niiiilatiinuila .Vlichener and Griswold. new lar to scutoaxillar suture (posterior margin of scutum somewhat species. smooth and shining but not forming broad shiny floor as in forms In Iiidniilhidium resembles a small with an open suture or sulcus) except suture foveate in E. (C.) na- appearance, (length sicum (Friese) ; posLspiracular fovea of propodeum much larger than 6.0-7.5 mm) Anthidium. largeh because the yellow metaso- 316 The UNrvERSiTi' of Kansas Scienc:e Bulletin

broadh concave but without lateral projecdons; S6 with mar- mal bands are dnidecl into four spots each, as in some species gin rounded, medially with rather dense row of the gin broadly margin of Anthidium. Unlike Anlhidium, depressed marginal of Tl to T6 lat- retrorsely cuned hairs. Posterior zones depressed of T5 is like the rest of the or a lit- dorsallv not different zone punctured tergum erally but scarcely so dorsally, punctation not differentiated about tle more densely so in the male, the zone from adjacent parts of terga, posterior impunctate margins as wide as base of first Piiiiilalioii: flead and tho- and its not angulate medially; the posterior margin flagellar segment. margin so that surface is dull; Indan- rax mosth finely and denseh punctate, punc- of T5 and other is rather broadly impunctate. terga tures of coarser and on shining surface; scutellum more in clvpeus thidium is similar to 'Pseudoauthidium, with which it agrees than coarsely punctate and more shining scutum; propodeimi sutures. It differs from that well the distincd> arcuate subantennal punctate throughout; metasoma with pimcuues separated a width; T4 to T7 in the immodified SI to S(> of the male, without combs by .shining ground, mostly less than puncture genus in size and un- wth more irregular and mixed or and S4 and S5 not especially concave. An punctures progressixeh' processes of outer surfaces of tibiae much coarser than as broad as shape; punctures usual feature is T7 of the male which is nearly in but those of thorax, hind tibial surface slightly irregular profile lateral latter short T6, \vith strong median and spines (the longest) not tuberculale. Pubesmue: Moderateh- long and jjallid except the median and lat- on dorsum of metasoma and almost absent on S4 and with irregular smaller teeth between and suberecl hairs on and dorsum of thorax. eral ones, .\nother distinctive feature is the maxillaiT palpus to S6; soiTie dirskv frons, vertex, as follows; basal two- Integument: Black with light yellow markings which consists of only one segment. thirds of mandible; clypeus; lower paraocular area, truncately This occurs in India. There is only one known on ver- genus ending at level of antennal socket; lateral trans\erse stripe on anterior of scu- species. tex; pronotal lobe; small lateral stripe margin anterior India Anlhidium. tum; axilla; medialh' broken posterior stripe on scutellunr, Etymology': plus on undersides and posterior .spots on tegula; longitudinal stripe of fore and mid femora, that on mid femur only on distal half; on outer side of fore Indanthidium crenulaticauda new species comparable distal area on hind femur; stripe and basal on outer sides of mid and (Figs. 3, 4, 9, 18-21) tibia; apical elongate spots hind tibiae; outer surfaces of basitarsi; comma-shaped submedian 6.0 head me- NL\le: Body length 7.5 mm, forewing length mm, marks on Tl to T5, these coalescing on T6 to form irregular width 3.0 mm. Head: Without carinae. Inner orbits converging dian blotch; transverse lateial marks on Tl to To; submedian and con- below. Clvpeus flat in profile, upper margin straight except lateral marks sometimes almost connected posteriorly. Wings with two teeth of vex laterally, lower lateral margin short, black, dusky, veins black. Small segments of tarsi brown; margins feebly conca\ e, rather thick, not over- sterna biownish; thiixl and mesalh , lower maigin simple, metasomal terga and following flagellar broadest hanging base of labrum. Labrum impunctate. shining, segments brownish black. bevond middle. addition at base" with strong erect lateral tooth slightly Fl.NLM.lC; Differs from description of male as follows (in so that one Mandible 3-toothecl but upper tooth ending obliqueh' to usual sexual cliaracters); Body length 6.0 mm, forewing length shallow be- with lower lateral could recognize four teeth with veiv emargination 5.2 mm, head width 2.7 mm. Head: Clvpeus outer surface with cari- refiexed narrow t\veen upper two teeth; coarsely punctate inargin lacking teeth, lower margin forming api- to base. Labial with first about half are fi\'e denticles. Mandible nae extending palpus segment cal shining ridge, beneath which suong of one al- lowest as long as second; maxillaiT palpus minute, consisting witli [\\e teeth, all separated by acute notches, interspace black with several bristles. (If this is a sec- than 2 to 4 which are teeth 2 most globular segment slightly broader interspaces equal; first is much broader than with ond segment, then the segment long to 5 subequal; oiUer suiface of mandible basally punctures in diT material.) Subantennal sutures dis- distance less than antennocular dis- and unrecognizable sparse. Interantennal slightly fVt)m to lower ends broader th;ui tinctly arcua'te outward, lines upper parallel, tance. Second and third flagellar segments much long, yen above tentorial Thorax: Hind basitarsus lower ends joining epistomal suture slightly segments four to nine all broader than long. distance scarcely than antennocular dis- toward all basitarsi shorter pits. Interantennal larger widest near base, tapering slighdv apex, to intcrocellar distance, than widths of ba- tance; ocelloccipiurl distance equal slightly than tibiae; basitarsal hairs mostly not longer narrower than e^e seen of hind tibia less than ocellocular distance; genal area sitarsi. Front and mid tibial spines strong, acute; apex of e\e. not level or Metasoma: T6 from side, widest near upper end Scape reaching with strong midapical blimt projection spine. ocellus; first almost denUculate carina, of anterior edge of anterior flagellar segment with strong, trans\ersc, preapical, depressed third broader than bv shin\ fo\ea twice as long as broad, second and slighth' long, medialh', aiid separated from apic;il maigin extending but tenUi as as broad, following segments slighth longer only long full width of tergum; S6 unmodified; impunctate tergal margins o\er 1 .5 times as as broad. Thorax: Without Punctation: and lather ele\enth slighdv long very narrow laterally. Clypeus coarsely on fineh and carinae except for strong carina, lamellate laterally, pronotal on upper half, more closely punctate progressively on lobe; omalus rounded; front end of scutum veiy gradually cuned densely so on lower half T4 to T6 with punctures finer than with down, without .smooth vertical surface; axilla and scutellum Tl to t3, less \ariable in size and shape than in male. Pubescence: from abo\e with feeble median scutellar of mandible margins rounded seen Scopa white. Integument: Basal third yellow. Clypeus and rounded as seen in that nearh meet cmarginadon, scarcely overhanging pro- black except for lower lateral areas preapically. without basal hind nbia file; scutoscutellar suture almost closed. Propodeum Mid tibia with continuous yellow suipe on outer suiface; fovea, vertical. middle of ubia; outer surface of series of pits and poslspiracular profile essenually with apical yellow area reaching sided, reduced Basitarsi about as long as remaining tarsal segments, parallel hind basitarsus with distal third black; lateral mark on T5 two to three mid basitarsus as long as tibia; basilars! with some hairs to trans\'erse streak; T6 black except for four small yellow spots of basitarsi. Front and mid tibiae each with times as long as widths forming a rectangle. with with median INDI.\; Poona: one small apical spine; hind tibia apex oblique Holotvpe male, 1 female and 2 male paratypes: ,\i-olia absent. Melasoma: in the Snow apical angle, tibial spurs nearly straight. l,onayla, Januan 20, 1959 (F. L. Wain), Entomolog- laterally, horizon- male is Tl with line margining basal concavity except ical Museum. University of Kansas, except one paratvpe enci- other to tal surface over half as long as vcrucal surface; tergal graduli transferred to the Natural Histon' Museum, London, the Tti with weak lateral T7 Utah State ing near spiracles, not bent back; angle; tlie USDA Bee Biolog^' and Sysl;emaucs Laboratoiy, transxerse, with about three-fourths as broad as T6. short, poste- University, Logan. Utah. shorter median and an notched or toothed, riorly directed lateral spine, spine, irreg- Et\'molog>':V,Vc»»to/»s, Latin, minuteh' and lateral SI to the male. ularly spinulose margin between median spines; raudaA.M\n. tail, with reference to T7 of mar- plus 84 unmodified, SI to S3 hairy and punctate; S5 with posterior Old W0R1.D AvrwD/ixF Bees 317

Larinostelis new geniis outer one strongly so. From and middle tibiae each with two short, apical spines, pallid and often difficult to see. Middle tibia Type species: Larinostelis scapulata Michciicr and C'.riswold, new species. nearly as wide as hind tibia, widest near apical third, anterior mar- gin strongly convex, posterior margin straight, .-\rolia absent. Tliis is a subgenus of small (6.5 mm long), robust, black MelasoiiKi. T2 widest; Tl \vith strong line margining basal concavity, horizontal surface somewhat shorter than vertical bees with extensive vellow markings. Like Slelis, the female surface; tergal graduli not bent posteriorly at sides: T6 transverse, not at all lacks a scopa. This is the most carinate of all .S/Ww-like bees, pointed apicalh . not denticulate, with strong, transverse, preapi- with carinae between the antennal juxtantennal present cal carina that appears to be the tergal apex when Tti and S6 are ax- sockets, and with carinae on the pronotal lobe, omalus, closed together: profile of T6 basal to carina short, straight; S6 with transverse basal or surface behind ca- illa, scutellum except median notch, basal zone of pro- strong, ridge carina, rina with many small tubercles, those near distal tergal margin podeum except medially and continuing behind spiracle, largest. Puiiclatioii: Coarse, especially so and cribrate on scutel- dorsal margin of metapleuron, longitudinally on hind ba- lum, axilla, and mesepisternum; elsewhere punctures mostiv sep- SI and across base of S6 of the fe- sitarsus, transversely on arated b\ aboiu half a puncture width: punctures fine in narrow male. The arolia are absent. The axilla is greatly produced space between juxtantennal carinae; tergal punctures coarse and rather close lateralh', fuier and more widely mid- laterally, so that it is behind the tegula. The two apical slighth separated dorsally; posterior zones of Tl to T5 slightly depressed, consist- spines of the fore and mid tibiae are miniUe. ing largely of impunctate, translucent marginal bands that are Larinostelis is known from a female from unique Ken)a. elevated abo\e surfaces of following terga. T6 with surface ante- It is not clear whether Larinostelis is a derivative of Stelis rior to carina coarsely, closely, and irregularly roughened. Pu- bescence: on dorsum of head and or an independently cleptoparasidc form. In the former case, Whitish, slightly dusky thorax, short, and surface onh on it would be best regarded as a subgenus of Stelis. The dis- sparse, inconspicuous, covering paraoc- nlar area: hairs rather dense on fore and mid basitarsi where largest covery of the male should help in deciding this question. hairs are slightly longer than width of basitarsus. Integument: Head with reference Etymology: Lariuos, Greek, fat, plus Stelis, and thorax black, metasoma red brown, except the following to the very robust body form. bright yellow: clypeus: lower paraocular area; small spot on ver- tex laterally; pronotal lobe in part: anterior spot on tegula; trans- verse mark lateralh' along anterior margin of scutum; axilla; Larinostelis scapulata new species narrow subapical, medially-interrupted band on scutellum; large (Figs. 22. 23) triangular mark on mesepisternum below pronotal lobe; ventral surfaces of fore and mid femora; tibiae except ventral surfaces; Fenl\le; 7.0 5.5 head fore lateral , Body length mm, forewing length mm, and mid basitarsi; large spot on Tl on T2 slight])' larger, width 2.6 mm. //rarf; Without carinae except foi" high, almost lamel- T3 with complete band; T4 to T6 yellow. The following red brown: that to late, juxtantennal carinae are straight, parallel one another mandible, scape, first two flagellar segments, tegula posteriorly, and separated by less than width of antennal socket. Inner orbits legs and terga except as marked with yellow, and SI to S6. Fla- below. in strongly converging Clvpeus convex profile, upper mar- gellar segments three to ten duskx brown. Wings dusk\', veins black. gin between subantennal sutures slightly arched, almost as long Holotvpe female: KENYA: 26 miles (42 km) southwest of as upper lateral margin; lower lateral margin less than half as long Nairobi, 5300 feet (1631 m), 1 1 Januan- 1970, M. E. Irwin and E. as any other margin: lower margin straight, denticulate medially. S. Ross, in the California .Academy of Sciences, San Francisco. teeth Mandible short, cui"\ed, 3-toothed, apices of acute, inter- Etymologi,': Scapulata, Latin, with shoulder blades, referring to spaces equal: outer surface shining, coarsely punctate, carinae the unusual lateral positions of the axillae. strong, not reaching base of mandible. Subantennal sutures nearly straight, longer than diameter of antennal socket, converging downward, suture well above tentorial In- joining epistomal pits. Genus Pachyanthidium Friese terantennal distance much less than antennocular distance; ocel- loccipital distance less than interocellar distance which equals to the of Pachyanthidium ocellocular distance: genal area about half as wide as e\e seen from Key Subgenera side, widest abo\e middle of eve. Scape not reaching ocelli; first 1 . carina absent and second flagellar segments subequal in length, much wider Eyes hairless; preoccipital laterally, than long; following segments progre.ssi\cly longer, seventh and behind eyes 2 about as as broad, ninth than broad, eighth long slightly longer —Eyes with abtmdant, short hairs; preoccipital carina tenth over 1.5 times as long as broad. 77(ora.v; With strong carina present lateralh , behind eyes (mandible of female on posterior lobe of pronotum and upper half of omalus, with 4-toothed) 3 lamella across upper end of metepisternum and across posterior margin of scutellum, the last denticulate; tegula widest at middle: 2. Arolia present; mandible of female 4-toothed axilla almost square seen from above, extending laterally behind (Namibia) Ausanthidiiim and in contact with scutosciuellar sutiue tegula (Fig. XXX): —Aiolia absent; mandible of female ustiallv 5- to 10- foveate, bottom of fovea shiny, fovea di\ided medially; scutellum toothed or dentictilate, -Ltoothed only in P. (P.) strongly produced posteriorly, strongly o\erhanging metanotum wicheneri Fasteeh s. str. and propodeum, profile acutely angulate, with strong median (Africa) Pachyanthidium emargination seen from abo\e. Pronotum with short, slanting, hair- 3. T3 to T5 each with slender lateral spine; ocelli less basal area, limited b\ trans\erse carina, and di\ided posteriorh- small, diameter about equal to width of base of first by numerous longitudinal carinae into small, lat- elongate pits, flasellar sesrment; hind coxa not carinate: arolia erally basal zone widening into large fovea delimited by carinae in male, absent in female (.Africa, southern above postspiracular fovea, also delimited by carinae; profile of present propodeum behind basal zone vertical. Basitarsi slender, nearly .•Vsia) Trichanthidium sided, about as as tarsal hind parallel long remaining .segments; —T3 to T5 without lateral spines: ocelli of ordinary basi tarsus with strong longitudinal carina on outer surface. Apex size, diameter greater than width of base of first fla- of hind tibia unmodified. Hind tibial spurs cuned near apices. 318 The Unix'Ersity of Kansas Science Bulletin

27 28

Figs. 22, 23. Larinoslflis smpulnia Michener and Gviswo\d, {em7i\e. 22. Face. 23. Dorsolateral view of head and tho- rax to show axilla (a) behind legula. Figs. 24-28. Pseudoanl.hidium (Tubemnlhidium) brachmtum Michener and Griswold. 24. l.aterofronlal view of male face showing mandibular dentition. 25. Same for female. 26. Apex of male nietasoma. 27. Dorsum of female with slip of paper under edge of axilla and side of scutcllum to show outline. 28. Forewing of female. Oi.li World AxminuM-: Bees 319

hind coxa arolia absent gellar segint-nt; carinate: odier, howe\'er, that they have frequently been given generic (possihhwitli wv\' minute arolia in both sexes) (east status. Cionuiion features, other than those inchcated in the

/\rabian Trichanlhidiodes to are 1 lack of carinae Mrica, Peninsula) key genera, ( ) except sometimes on the pronotal lobe and sometimes on the preoccipital ridge behind the vertex, (2) the subantennal siuures aris- Subgenus Ausanthidhim Pasteels straight at ing or near the tentorial pits, (3) rounded scutelhun not or little the in There is a single species, Pachyanthidium (Ausaulhidniiii) ON'erhanging luetanotum, profile rounded or ausense (Ma\Tomoustakis), new combination. in the subgenus SpinanthidieUum, angled, (4) presence of the of of This subgenus has not hitherto been placed in Pniii\a»- arolia, (5) tendency T6 the male to be trifid or trilobed in the thidium, but nearh all its characters support such a placement. (not subgenus SpinanthidieUum), ((i) the small, biU little-exserted male T7 which is 3-toothed or in A small ft)\ea behind the propodeal spiracle, reported by Pas- the Pksianthidiums. str. and Carinanlhidium'whh teels , that ( 1969a) might support the original placement in subgenera is ill the middle tooth reduced to a minor between two AnlhidieUum, defined, no wider than the spiracle itself, convexity sterna and probably merely the edge of the spiracular area, not a long, widely separated teeth, (7) male litde modified 85 with absent in the fovea in the sense of that of Aulhid/c/liuii and other genera. except apical comb, subgenus Spinau- thidium, and S(i characteristically lobed or with various con- vexities, (8) male and Subgenus Pachyanthidium Friese s. str. gonostyli simple, slightly expanded hairv apically, with two small teeth at apex in Plesianthidium s. str. Of the four A remarkable feature of the subgenus is the \ariabilit\ in subgenera, two, Ph'sianthidium and Spinan- dentition of the female mandibles, from 4-toothed to mul- thidium. ate closely related and might well be united. titoothed and to minutely serrate. Males also are variable in mandibular dentition, from 4-toothed to 8-toothed. In Key to the Subgenera oi Plesianthidium spite of this variation, the species are similar in most of their characters and constitute a morphologicalh rather ho- 1. Preoccipital ridge with carina behind vertex; T(i mogeneous taxon. of male truncate, without lateral tooth, distally with longitudinal ridge, highest at posterior margin of Trichanthidiodes new subgenus tergum; mandible less than twice as long as broad (Cape Province) SpinanthidieUum Type species: Pritliyiinlliiiliiim semdutnim Pasteels, 1984. —Preoccipital ridge not carinate; T6 of male with me- dian truncate or roimded to Presumably because of its hairy eyes, P. scmiluteiiin was pointed projection, and lateral tooth; placed in Trichanthidiitmhy Fzsteeh (1984). It differs, how- rarely weakly produced, strong mandible over twice as long as broad 2 ever, in many ways including its extensive yellow coloration. 2. S6 of female with median ca- The following are subgeneric characters; in those marked strong longitudinal rina; hind trochanter of male with mesal (1) Trichitulhididdcs agrees with Pachyanlhidium s. str., in subapical almost all black white on those marked (2) it agrees with Ausanlhidiiim: and in those spine; pubescence except face of male (South Africa) Carinanthidium marked (3) it agrees with Trichanthidium. —S6 of female not hind trochanter of male Eyes hain' (3): ocelli not reduced in .size (1,2); niaiidible of fe- carinate; 4-toollied male (2, 3) ; omalar carina continued direclh' acro.ss ven- not spined; pubescence brown to gray, or whitish on tral midline (3); wing.s not strongly darkened (2); arolia absent in face and venter 3 male (1); arolia absent in female (1, 3); hind coxa carinate ; Tl 3. T7 of male trifid, median tooth or lobe with strongly lransver.se carina not double laterally, i.e., vvithoiil preapical lateral ones; S5 of male without carina with 1 exceeding apical (1,2); terga wide, transparent, inipunclatc margins ( , comb; ; without lateral with lateral 1 2) terga spines, rounded .swellings ( , miudllaiy palpus 2-segmented (Cape Province) male wilhoiil 2);S4of comb (1, 2) ;S5 of male with comb (l);gonos- Spinanthidium of male with club tylus delimited basallv b) strong carina (3); T6 —T7 of male with median tooth reduced to low of promi- female with small, deep midapical emargination (1, 2); S5 of nence so that is essentialh' bifid; S5 of male female flat, not biconvex or bilobed (1.2). tergum with comb; maxillan' This subgenus is known from Kenya and Saudi Aiabia. apical palpus 3-segmented (Cape Province) Plesianthidium s. str. There is one species, P. (Trirhaiilhidiodi'.s) semilulcum Pasteels. Etvmolog)': Trichanthidium plus the Greek suffix -odes, sim- ilar to or resembling. Genus Pseiidoanthidium Friese

Pseudoanthidium is Genus Plesianthidium Cameron here used in a broad sense to include ninuerous species having compact bodies, commonh' with Plniaiilhidittiii consists of four in subgenera that agree the the form of Anthidium s. str.; except in the subgenus Mi- lack of pale markings except on the face of the male and cranthidium the head is thick and the tibiae are tuberculate in other characters indicated in the key to genera and on the outer surfaces. Even in Micranthidium the tibiae are below. The subgenera are different enough from one an- coarsely punctate. 320 The UNi\'ERsrr^()K Kws.vs Science Bl'i.letin

Key to the Subgenera oi Pseudoaulhidinw in part on identification of Antliiiliinii aljiiiiumMoramtz, the type species of Psi'iidoanlhidiuni. V\'e have seen specimens of various 1 . identified as One of them Gena margined posteriorly by distinct preoccipi- species alpiiium. agrees with the a black tal carina (gena narrower than eye seen from side) original description, having clypeus in the 9 male, an imusual feature, and we believe it is true alpinum. All of in —Gena without preoccipital carina (sciitelliim them, however, agree subgeneric characters with Parauthidiclhnii which we therefore as a roimded or medially emarginate seen from above; place synonym of tibiae tuberciilate on outer surfaces at least in female; Pseudoauthidiums. sU". Pasteels (1969a) and Warncke (1980), Anlhidium barbatiim and male without lamellate laleial lobe on S3; clypeus however, place Mocsarv' setraticeps Friese in the of A. This cannot be cor- of female with discal hairs straight) 3 synonymy alpinum. for the of in Berlin is a robust 2. Preoccipital carina behind \'ertex strongly pro- rect, type senaticeps larger, of the it is a female with a duced back over front of thorax as a lamella; pos- species subgenus Royanlliidium; terior basi tarsus of male more than twice as long as bare, projecting dentate clypeal margin. Morawitz said that broad (scutellimi broadlv triuicate in dorsal view, A. alpinum (based on one male) is a species with the aspect of Pseudoanthidium lituratum is posterior margin angulate laterallv near axilla; tib- (Panzer), which a Pseudoan- thidium s. str. as here understood. As to it con- iae not tuberculate but coarsely punctate; lateral lobe Carinellum, tains two and its distinctive which of S3 of male lamellate; clypeus of female with dis- only species characters, cal hairs bent clown (Africa, .Aiabian Peninsula) appear to be apomorphic relative to other Pseudoanthid- — Miminthidiuin ium, do not seem to warrant a separate subgenus. Preoccipital ridge behind vertex rather weakly car- inate; posterior basitarsus of male less than twice as Subgenus Royanthidium Pasteels long as broad (west Africa)

Semicarinella As indicated in the above list, we consider Reanthidium Pas- 3. Propodeum shagreened; TG of male with strong teels to be a synonym of Royanthidium. Reanthidium differs which is fore apical flange medially emarginate; from Royanthidium in having the preoccipital ridge cari- and mid basi tarsi of female with hair long plumose nate behind the \'ertex. We judge that a separate subgeneric and southern Tuberanthidiuiii (Tanzania Africa) name is not needed for the one species with this carina. /-". — where not T6 of male Propodeum shiny punctate: (R.) nigiicolle (.Vlorawitz) , placed in Reanthidium h\ Pasteels, without fore and mid basitarsi emarsjinate flansre; is a synonym of P. (R.) melanunim (Wug), the type species of female with hair not 4 long plumose of Royanthidium, according to Warncke (1980). 4. T6 of female with surface broadh- and conspicu- ously excavated; T5 of female with small, midapical Subgenus Semicarinella Pasteels marginal projecting lobe; exposed part of T7 of male than of longer exposed part T6, vei")' deeply Semiearinella is based on a single male specimen that we bilobed; S5 of male without strong lateral tooth or have not seen. It may be merely a species of Mieranthidium lobe basin to cend"al Asia (Meditenanean and Sudan) with unusual characters. — Exiinlhidium T6 of female with surface largely convex; T5 of Tuberanthidium Pasteels female without midapical lobe; exposed part of T7 Subgenus of male shorter than part of T6, exposed weakly Pasteels considered that an iniusual featiue ol ilu- males to strongly bilobed; S5 of male with strong lateral of Tuberanthidium \va.s the 5-toothed mandible. Most speci- process 5 mens ha\e four teeth; often the number of teeth on opposing 5. Clypeal apex of female fulK' exposed, coarsely den- mandibles differs; in one specimen one mandible has only ticulate, proUuding; S3 and S4 of male broadly emar- three teeth. The definition of this subgenus has been con- ginate or S3 with median V-shaped emarginarion and siderabh modified to include the following new species. S5 with apicolateral projection, all with< )ut combs or specialized hairs (Palearctic) Royanthidium — Pseudoanthidium (Tuberanthidium) brachiatum new species Clypeal apex of female largely hidden by hair, with- (Figs. ."), (i, 'i4-2S) out shining, protruding premarginal ridge; S3 of male with comb of long, wavy bristles and S5 with This species differs in several major characters from other comb at apex of apicolateral projection species of Tidxraiilhidiiim: our initial \iew was that it slioiild Ix' placed in Pseudoanthidium s. str. a new subgenus. However, we decided that another monot\ pic subgenus is iiniiecessan' and ha\e broadened the definition of I'li- beranthidium instead. It differs from other species of Tuberantlnd- Subgenus Pseudoanthidium Friese s. str. ium in the more angulate (but not carinate) omalus, lack of a dor-sal bulge on the mandible of the female, gently convex apical mar- The oi Pseudoanthidium Pamnthidiellum in- synonymy and gin of S3 of the male with wa\y bristles (on midapical part of ster- dicated in the List of Genus-Group Names above is based num) inconspicuous and shorter than white hairs located more Oi.n VVoRin A.vni//)ii.\r Bees 321

and black laterally, combs on apical pan of apicolateral Pubescence: projection ground. Spmse except long, plumose, dense, and white of S5 of the male and of small combs on the of on pair margin S5 on apical clypeal margin, thoracic \en'ter, and SI to S3; either side of the pubescence midline. The structure of S5 is similar to that of ol vertex, dorsum of thorax, and metasomal terga dusky except various other o[ Pmidonnlhidium \ellowish on subgenera and reinforces our \ iew posterioriv meta.soma; othenvise pubescence white that Tuhemnthidium s,\\ou\A or on side of thorax and be included in Psnidoanlhidium. on legs, yellowish. Inlci^umi'iil: Black with M.AL1-;: Boch 8.0 to 10.0 \ellow as follows: mandible black length mm; forewing length 7.2 to 8.0 markings except teeth and api- head width .S.8 to 4.2 cal and inner mm; mm. Head: Without' carinae. Inner ot-- margins: civpeus; lower suprachpeal area to about bits below. level ol antennal sockets; lower converging Clypeus flat in profile, upper margin paraocular area up to same level, next to strongly arched; lower lateral margin xeiT short, lower margin slightly higher socket: transverse mark laterally on vertex- spot on front of straight, apparenth- slighih imdulate, completelv covered bv long. tegula; legs except coxae, trochanters, and femora white, downward basalh; broad transverse bands on Tl to directed, plumose hail ; chpeus not overhang- T5, broadiv interrupted base of labrum. Labrum widest at on Tl, less .so on ing base, otherwise parallel progres,sively succeeding terga to incompletely sided, with or not so on T5: T6, T7, and sterna longitudinal depression between midlateral ridges. entirefy vel'low. Posterioi- mar- Mandible 4-toothed, of gins of Tl to Tfi translucent brown apices teeth acute; interspaces equal and grading to translucent vellow acute lower on notches; and upper teeth larger than two median posterior terga. Wings duskv, xei'ns black. teeth; outer surface of mandible FiiM.\i,i-;: Differs from of as largely smooth, shining, sparseh description male follows (in addition without carinae. to usuaUexual Both punctate, Second segment of labial palpus oxer characters): length 8.5 to 9.0 mm; forewing 1.5 times of first; 7.0 mm: head \\idth 3.9 length maxillaiy palp\is short, 2-segmeiued. Siil> length mm. Head: Lower margin of chpeus antennal sutures with three denticles, hidden as in strongh' arcuate outtvai d, lines honi'upper to lower male. Mandible .5-tooilied, upper ends k)wei- ends tooth parallel, joining epistomal suture much abo\e ten- largest, obliquely truncate, thus suggestive of additional torial tooth pits. Interantennal distance over 1.5 times antennocular (Fig. 25). Interantennal distance litde greater than anten- distance; nocular ocelloccipital distance greater than ocellocular distance, distance; ocelloccipital distance equal' to or less than ocel- which is locular which is greater than interocellar distance; genal area nearh as distance, greater than interocellar distance. All wide as seen eye from side, widest behind upper third of eve. flagellar segments except first and last (which are about as in not male) broader than Scape reaching level of anterior margin of anterior ocellus, long. Thorax: fore femur not angulate at base. first 1 .5 Hind basitarsus widest near fiagellar segment times as long as wide, second wider than base, tapering: long hairs of basitarsi plumose, of tibiae oii long, subsequent segments progressively longer, tenth about as long .\pical spines sharp, large fore and mid tib- as broad and iae. Mrlasima: eleventh over 1 .5 times as long as broad. Thorax: With- Lateral longitudinal carina of T5 as in male. T6 with out carinae one on flat for except pronotal lobe; upper half of omalus profile except strongly elevated apical lamella across area but not about halfas wide as basal strongly angular carinate; .scuUim with anterior end cuned (exposed) width of tergum, the lamella strongly down but without \enical siuface; axilla and scutelhm-i produced apically to form projection on each s'lde of small me- dian trans\erse, margins rounded as seen from alxne with ^veak median notch; laterallv, near end of lamella, it forms small shoulder nearer acute as seen median notch than lateral ,S(i emargination, posterior edges in profile, o\erhanging extremitvofTfi. unmodified. nietanotuni and Punclalion: le.ss propodeum; scuto.scutellar .suture narrowly foveate. Clypeus shining, coarsely and closeh" punctate ex- of scutum for which is posterior edge impunctate, supplementing width of foxea. cept uppermost margin smooth, like lower margin of without basal area. Outer suiiiices of Propodeum series of pits and postspiracular fo\ea. su]3racl\7X-al ubiae more strongh tuberculate Front femur thick at than in males. Middoisal of Tl to base, forming ventral angle at ba.se. Basilars! parts T4 with punctures mostly as as tarsal h\ se\eral widths. long remaining .segments together, parallel sided, all separated puncture Pubescence: Scopa yellowish white. shorter than tibiae, fiont and mid basitarsi with some hairs aboiu Hairs of coxae, trochanters, and bases of femora hooked twice as as basitarsal or somewhat wa\y'. long diameter. Front and middle tibiae each Integument: Suprachpeal area \ellow onlv along with one small, blunt lower toward antennal apical spine; hind tibia with apex oblique margin, extending up base laterallv; small with median tibial segments of tarsi yellowish brown; vellow band of apical convexity; spurs slightly cuned at apices. T5 narrowly in- Ar()lia absent. Mrlasoma: Tl and of femora and meta.somal T2 widest'; line margining con- terrupted: parts sterna \ellow brown. of Tl male, 1 male and cavity distinct, extending to side of tergum, horizontal sur- Holotype 2 female paraivpes: TANZANIA: face of Tl over half as National Park, 2300 long as vertical surface"; graduli of Tl to T4 Tarangiri feet (708 m) altitude, 23 January near on T5 1970 (M. E. Invin and E. S. The ending spiracles, cumng postenorlv and forming Ross). holotvpe and female carina are in the California of strong reaching posterior tergal margin ancl forming small paratyjses Academy Sciences, San Francisco; lateral lobe or ver\ blunt tooth; Tfi the male in the USDA Bee without lateral tooth, with pro- paratx pe, Biology and Systemafics Lab- lamella that is absent in oraton. Utah State Utah, jecting apical middle third, leaving de- University, Logan, flanked Brachialus. Latinized Greek pressed emargination by highest part of apical lamella: T7 Etymology: meaning with arms, re- to the small, expo.sed part one-third as wide -as T6, bilobed with semi- ferring armlike lateral proces.ses of S5 of the male. cncular between emargination lobes; SI to S3 simple with abun- dant hair; S.'? with xellowish long wavy bristles arising near margin ol middle third, these bristles inconspicuous, their'cunature not Genus Rhodaiifhiditim Isensee hairs lateral to them strong, while and longer than waw bristles; S4 with and margin broadly distinctly concave; S5 margin also con- Ke\- to the Subgenera oi Rhnddiilhidiuni cave, with small \eiy black comb on each side of midliiie, produced at side to slender, lateral long, arm directed each 1 . Tfi of male with posteromesalh, median, produced ti iincati( m or with black comb of six apical or seven teeth; SO with broad me- rounded and dian, process lateral tooth or shoul- apically notched projection. Posterior zones of Tl to T5 strong der; S5 of male with lateral onh' medialh', distinctly so strong tooth and median slightly depressed laterallv, more finely than rest of comb; omalus ])unctate terga, nearly half of each zone forming marginal weakly carinate above; mar- broad impunctate elevated abo\'e base of of .scutellum with margin following tergimi. gin sharp edge laterally; female I'linilaliuii: Rather dense, finer on frons and vertex than on tho- with apical projection of fore and niid tibiae n;irrowlv rax, with of clypeus shining punctures upper part well separated bidentate and shallow, lower of (western Palearctic) Rhoddnlhidium s. str. margin siipraclvpeal area impunctate, shin- —Tfi of male or with ing; mesepisternum with punctures contiguous; propodeum simple scarcely produced broad strongly shagreencd, with punctures onlv on basal and lateral truncation, with or without strong shoulder or weak areas; outer surfaces of tibiae coarsely punctate and moderately lateral tooth; S5 of male without lateral tooth, with- tuberculate; meta.soma! punctures well bv separated shining out comb; omalus not carinate or xveaklv so above 322 The UNHTRsm- of Kvnsas Sc:ienc:e Bl'lletin

in its rel- R. {Asiauthidium} cnturigejise (Ciiraud) and its robust body, well-developed vertex, broad middle tibia, atives; margin of sciitelkim rounded, not forming and the often oblique \ein cu-v of the hind wing with the sharp edge except in K (Asianlhidium) caturigense; fe- second abscissa of M+Cu shorter than in other Stelis al- male with apical projection of fore and mid tibiae though not as short as in most Trachusa. However, Protostelis either not notched or widely and shallowly emar- does not have the reflexed male T7 characteristic of Tra- ginate 2 chusa and does have Stelis-Wke features such as the Uvo spines 2. Trifid apex (median tooth sometimes broad and on the apices of the front and middle tibiae and apically ex- rounded or trimcate) of T7 of male occupying full panded, angulate male gonostyli. We therefore regard Pro- width of tergum; pronotal lobe with lamella or ca- tostelis as a subgenus of Stelis, not related to Trachusa, but weak in R. lAsirnithiflnuii) either the S'/fV/s-like or the Trachusa-Yxke rina, glasunoiii (Morawitz) ; characters might re- head with backgroimd color black (Mediterranean suit from convergence. basin to central Asia) Asianlhidium —Trifid apex of T7 of male much narrower than ter- Key to Old World Subgenera of Stelis gum, which has lateral angles so that tergum is 5- toothed: lobe withoiu or with xeiT weak pronotal 1 . Clypeus produced well over mandibles and apex carina; head to central largely yellow (Tmkey Asia) bilobed, strongly so in female; anterior spine of Megaulhidiiim fi'ont and middle tibiae conspicuous, enlarged, curved posteriorly (Palearctic. to Kenya) Stelidomorpha — not over mandibles, trim- Subgenus Asianthidium Popov Clypeus greath' produced cate or subtruncate; anterior spine of front and The synonymy indicated in the list above is based on sim- middle tibiae less than twice the size of posterior 2 ilarity and the small munber of species involved. Asianlhid- spine ium in its original narrcjw sense, Axillanthidium, and 2. Axilla projecting laterally beyond lateral margin of Oxyanthidium each contains only a single species. Asianthid- sciumn; sciuellmn strongly projecting over meta- ium in the sense of R. (A.) glasunovi (Morawitz) seems the notum and propodeum (southeast Asia) . .Malanthidium — not most deserving of separate subgeneric recognition if the sub- Axilla projecdng laterally beyond margin of scu- genus as here imderstood is to be di\ided. Its weak rather tmn: scutelliun at most weakly projecting over meta- than strong carina on the pronotal lobe and the elexated notum and propodeiun 3 3. carinate median process of T7 of the male (suggesting Rliodanthid- SciUellum not laterally; head and thorax iums. str.) differentiate it fiom other species. K (A.) anikatum witholU light markings; hind tibial apex with two (King), the species placed in Oxyanthidium. differs from spines or angles, one near outer middle of apical tib-

ial with .S'. other species in the sDongly denticulate and laterallv angulate margin (if only one spine, simiUima margin of T6 of the female and the translucent preapical Morawitz, not on posterior apical angle) or the lamella above the apex of S6 of the female, so that the ster- spines united to form trimcate margin, the area num has a double margin. sparsely hairy so that structure is easily seen; oma- Placement oi Axillanthuiium in the syncjnvmy of As/V/z^/Z^/V/- lus not carinate (Holarctic, Oriental) Stelis s. str. ium is based on Warncke (1980), who considered Axillan- —Scutelhun carinate laterally; head and thorax with hind tibial with a thidium axilkireVasteels as a synonym oi"Anthidium caturigense light markings; apex single spine a mere hidden in hairs rf)traZf Morawitz" and on a specimen of caturigense, the type (sometimes angle) largely species of Trianthidium, with axillae emarginate and ante- near the posterior apical angle of the tibia, in front riorlv angulate, nearlv as accentuated as in Pasteels' illus- of which the apex of the tibia is a convex margin; carinate 4 tration of axiUare. Other specimens of caturigense have the omalus usually emargination and anterior angle scarcely evident. 4. Hind basitarsus with carina along inner dorsal angle; mid tibia flattened, apically enlarged, twice as wide apically as basally; SI with transverse carina Subgenus Rliodanthidiiim Isensee s. str. overhanging apical margin (Holarctic) Protostelis —Hind basitarsus without carina; mid tibia not flat- Most of the species form a imified group, but R. (R.) in- tened, little enlarged apically, apex at most 1 .5 dmes fuscatum (Erichson) is quite clistincti\e and has been given basal width; SI without transverse carina (Palearc- the genus name Bellanthidium Pasteels. Since there is only tic) Pseudostelis one such species and its relationship to the others is clear, a subgenus for it seems unnecessary. Subgenus Malanthidium Pasteels Genus Stelis Panzer Anthidium inalacccnse Friese, the type species of Malan- " thidium and known from the male, was marked "?Stelis The suggestion has been made that the subgenus Protn- onl)' in the its transfer to Stelisis therefore stelis might be derived from Trachusa, and that Stelis there- original description; not entirelv We ha\e seen a female of a related fore might be polyphyletic. /'roto5te/w resembles Trachusa in surprising. Oil) World Amiiidiixe Bees 323

that lacks species a and is a Slelis 1. scopa obx-iously s. Pas- Key to the Old World of Trachusa teels Subgenera (1969a) described Stelis malaccemis (Friese) (as Malan- thidium) (new combination) in some detail. His 1 . Mandible of female with three figure 138 more or less equidis- shows a tant midapical chpeal denticle, also mentioned in liis teeth: maxillan palpus as long as maximum description. No such denticle is on the width of present t\'pe (and galea, 4-segmented (yellow markings absent for face of only known?) specimen although there is a small shinv except male) (western Palearctic) on the place clypeal margin. Trachusa s. str. —Mandible of female 4- to 7-toothed; maxillan pal- pus shorter than width of 3- or Subgenus Protostelis Friese galea, 4-segmented 9 2. Second recurrent vein The placement of Doxaulhidiinii in the s\non\iii\ oi Piu- entering second submar- lostelis'm the above ginal cell basal to second transverse cubital List of Geniis-Gronp Names is based on \ein; T7 of male with median basal descriptions and figiues (see Pasteels, 1969a), on Warncke projection (Mediter- ranean basin to central .' (1992) and on a personal communication from G. van der Asia) \rchtanthidium —Second recurrent \ein Zanden . or distal (1993) Friese (1911) also suggested that Anthidinm meeting to second transverse cubital vein; T7 of male paradoxum Mocsarv might be a Protostelis and Warncke without basal pro- (19S0) placed it in .SV^'fo. jection 3 Two one 3. Subantennal suture arcuate groups, composed of larger species, the other distinctly outward; of have of male bifid, smaller, commonly been included in Protostelis. Pas- gonoforceps deeply Y-shaped (male teels indicated that unknown in Onhanthidium) 4 (1969a) the laiger and smaller groups (Palearctic) be —Subantennal suture of might independent and thus separate subgenera. We nearly straight; gonoforceps the male not place larger species in the subgenus ProtosU'lis and the Y-shaped (not palearctic) 5 smaller 4. Scutellum tiimcate as seen ones in the subgenus Pseudostelis. The size differences posteriorly from above, are not consistent, bin in margin cuned sharply forward to meet general apply; S. signata (La- laterally is of axilla; of treille) often onlv b mm ,S'. strandi is longitudinal margin eyes female con- long, 6 to 8 mm long, while the larger species measure 10 mm or more. Both spicuously diverging below; interocellar distance of female less differ from other Stelis in the much than half distance groups 3-segmented maxillaiy ocelloccipital palpi and the combination of scutellum (southeast China, Taiwan) Orthanthidium carinate laterally but —Scutellum rounded not strongly overhanging the propodeum. Both ha\e broadly except medially emar- on the ginate; lateral of axilla convex but in light markings head and thorax and the omalus f re- margin gen- eral carinate. The slanting; eyes of female interocellar quenth- anterior suiface of the mesepisternum subparallel; is distance about half of always concave. Protostelis differs from Pseudostelis and ocelloccipital distance other Stelis in several In (Mediterranean basin to southeast China, ways. Protostelis the apical angle of Oriental) the hind tibia is dorsal rather than along the outer face; the Paraanthidium hind basitarsus has a 5. Omalus distincth' carinate, carina onto longiuidinal carina along the inner dor- extending vential surface of thor^LX sal the mid tibia is close to middle coxa (.Africa) margin; flattened, much enlarged api- and in the cally female is clothed with dense decumbent hair; Massauthidium and the —Oinalus not carinate apical width of the female mandible is 1.5 to 2.0 g times the narrowest width. 6. Arolia present in female, absent in male; 84 and Protostelisk fiuther distinguished S5 of male without from Pseudostelis bv the verte.x, wide combs (southeast Asia) long impunctate api- cal and the Metatrachusa tergal bands, strongly carinate preapical mar- —Arolia if gin of SI. minute present; S4 and 85 of male with combs as in Paraanthidium (west .Africa) .Congotrachusa Geniis Trachusa Panzer Subgenus Archianthidium Mavromoustakis Several characters that ate often stable within a genus vary m Trachusa. Aj-olia are commonly present but sometimes Although this subgenus was not included in Trachusa reduced or by greath- absent. Combs on S4 and S.'i of the Pasteels we believe (1969a), that it easily falls within that male are sometimes but also often present absent, and genus. Superficially, because of its size (12.5-18.0 mm nvAx long), van even within a (of the Heterau- form, and abundant species subgenus yellow markings, it resembles the North see thidium, Brooks and Griswold, 1988). The number of .American Heteranthidium subgenus and well-marked species mandibular teeth of the female is of commonly four but may Paraanthidium. although unlike the former, the be three yellow as in Trachusa s. str. or fi\e, six or seven, as in the metasomal bands at least on Tl are broken It is Massauthidium and medially. subgenera Congotrachusa. The number indeed similar to and related to of probably Heteranthidium. segments in the maxillan ma\' be four or re- as indicated the paljDus especially by large, volsellalike process from duced to three. the of the male apex gonocoxite (genitalia of three species illustrated by Mavromoustakis,, 1939). .All of its characters 324 The UNr^TRSiTi- of K\.\sas Science Bl'ixetin fall within the range of variation for Hetcrauthidhnn except Trachusoides simplex new species the broken metasonial bands, the position of the second re- (Figs. 29-33) current vein (see ke)' to subgenera), and the midbasal, Structural characters that differ from those oi .\j}ianthidium are retrorse, blunt or truncate process of T7 of the male. (T7 italicized. does not have two slender apical processes as suggested by Fknl\lp:: Bod) length 12.0 mm, forewing length 8.0 mm, head there are width 3.9 mm. Head: Carinae absent, hiner orbits Pasteels, 1969a, fig. 12. Instead nvo strong carinae subparallel, upper halves genth concave. Clvpeus convex in profile, hexago- on a high ridge extending from the basal process toward nal, upper margin between subantennal sutures arched, lower the of the .Ajolia are in both sexes apex tergum.) present margin with eight denticles (lateral ones weak) between which arise the often does not show four and female mandible recog- tufts of bristles (median ones longest) . Labrum rather weaklv scle- nizable teeth, the upper two being united in a sometimes rotized, little longer than broad, broadest at ba.se and at apical fourth, surface minutelv with tufrs of erect bristles undulate margin. punctate, apex (Fig. 29). Mandible rather long, 4-toothed (Fig. 29), outer surface dull and minuteh roughened except for smoother distal margin and apex of rutelUun. outer and condvlar ridges ven narrow but Subgenus Paraanthidhim Friese stronglv elevated, shining, outer ridge extending from cap of rutellum about halfwav to base of mandible, cond\lar ridge ex- Megachile steloides Bingham, 1896, the type species of tending somewhat farther. First segment of labial p;ilpus about twice Protanthidium, a oi Paraanthidium, when junior synonym as long as second; ma-\illan palpus small, .3-scgmented, second seg- transferred to Anthidium, became ajunior secondary homo- ment much longer than first, third minute. Subantennal sutures re- than diameter antennal socket. downward, nym oi: Aiilhidiiim sti'loidt's Spino\-a., 1851. It was therefore stiaight, longer of con\erging suture abo\e tentorial hiterantennal distance named as Anthidium lonsicurnehy Friese, 1902. Even thousrh joining epistomal pits, about twice antennocular distance; head well de\eloped behind the two are no in the same Friese's species longer genus, and aboN'e ocelli so that ocelloccipital distance is nearly twice interocel- name is retained, as Tradiusa longicomis (Friese), according lar distance; ocellocular distance about 1.5 times inlerocellar distance: oceWi area almut as to Article 59(b) of the International Code of Zoological largely below upper ocular tangent; genal as wide eye Nomenclatiue. seen from side, widest near middle oj eye. Scape reaching about le\el of middle of anterior ocellus; first flagellar segment 1.5 times as Orthanlludium, known only from females, could be con- long as broad, second and third broader than long, fourth nearlv sidered as a of Paraanthidium. If the unknown synonym as long as hxoA.A, foUoieing segments progressively longer until ninth is male of Orthanthidiiim has the characteristic features of about 1.5 times as long as broad: tenth o\er twice as long as broad. Paraanthidium, the two subgenera should be united. Thorax: Without caiinae except for strong canna on pronotal lol>e: oma- lus rounded; front end of scutum gradualK bent downward, with- out smooth vertical surface; axilla and scutellmn with posterior convex seen from above for small median scutel- Trachusoides new genus margins except lar emargination, these margins not carinate, rounded in profile, metanotinn and base of scu- Type species: Tirirliiisoides sniiptex Micliener and Ciriswold, new species. stronglv overhanging propodeum; toscutellar siUure closed, not forming fovea. Propodeum with no indication of biisal or fo\ea, con\ex, This genus contains a large species (length 12 mm) sim- ]5its post-spiraculai' profile upper two-fifths being declivous but not vertical, cuning gradually onto ilar in form to Apianthidium and Trachusa. The head of the \erucal lower three-fifdis. Basitarsi each longer tlian remaining tivrsal female is black, the thorax has limited pale yellow markings, segments together, widest near bases and slightly tapering; mid and the metasoma is yellowish red. A unique character is basitarsus as long as tibia; hind basitarsus o\er three times as long as broad. Front and mid tibiae each with mid- the simple female claws, breaking down an otherwise con- single, strong, blunt, hidden bv short \vmy\ mid tibia stant character of cleft or toothed claws in female Anthidi- apical spine largelv pale Jor shape of see generic diagnosis: hind tibia with apex truncate, apical margin ini. (The inner tooth is veiT small, however, in Serapiita.) onh' slightlv con\ex medialK: tibicd apurs nearly stmight. C.lcnos sim- for a carina on the there are Except strong pronotal lobe, ple. Arolia absent. Metasoma:T\ to T3 widest; Tl with carina mar- no carinae on the head and thorax. Basal or postspiracular gining basal concavitv, horizontal surface less than half as long as vertical surface; tergal ending near spiracles, not bent pits or foveae on the prcjpodeum are absent. The middle graduli posteriorlv; T(i unmodified, profile straight, margin not denticu- tibia (female) is narrower than the hind tibia, with anterior late; S6 slighd\ exceeding T6 medialh', not modified. Posterior zones and and s\nimetricall\' convex seen posteiior margins equally ofterga scarcelv depressed; no conspicuoirs impunctate margins. laterally, the apex as narrow as the base of the tibia; these Punctation: Dense throughout, finer on cl\ peirs and lower paraoc- tibial characters differentiate it from both Trachusa and ular and genal areas than rest of head and thorax; propodeum punc- tate throughout; tibiae more coarseh pimctate than body but Apianthidium. The male is imknown. surfaces not tuberculate; posterior depressed zones of Tl to T5 Trachusoides is found in southern India and known from and all of T6 more fineh' punctate than clypeus; sterna with basal a of T. Michener and Griswold. single specimen simplex and apical zones not or sparselv punctate. Pubescence: Pallid but The only known specimen of Trachusoides is described somewhat vellowish on head and thorax, dusky on upper part of head, more on metasoma and Hairs rather fulh below. It is similar in manv features to Api- orange legs; scopa orange. of clvpeus mostly erect and ven short, scattered longer hairs anthidium. When the male is known, it may prove to be a preapicallv; bristles of clvpeal and labral apices orange; lower mar- of that in of the claws of the fe- species genus spite simple gin of mandible with series of about six widelv spaced long, erect male, which are unique among Aiithidiini. bristles (as in .\pianthidium) . Rest of head and thorax with hairs rather on sides and venter of thorax and on Etymology: Trachusa plus "-oides," similar to. sparse, longest propodemn. Metasoma with hairs short and suberect dorsally, quite dense, denser and directed posteriorly on posterior tergal Old World Anthidiink Bees 325

^0 i^r L^i

32

Figs. 29-33. Trachusoides simplex Michener and Griswold, female. 29. Slightly laieral \icw of face to show mandibii- lai' dentition. 30. Slightlv lateral view of thoracic dorsum. 31. Middle tibia and basitarsus, outer view. 32. Forewing. 33. Claws of middle leg.

zones and T6. .Scopa long and dense, including that on S6. Iii- Etv'mologv': Simplex, Latin, simple, with reference to the simple tegumenl: Black on head and thorax except dull pallid yellow on claws. broad lateral margin of scutum, almost whole axilla, and poste- rior two-thirds of .scutellum. Second and third llagellar segments, part of pronotal lobe and preaxilla oiange brown. Tegula, tarsi, LITERATURE CITED tibiae, and apices of femoia orange. Wings strongly dusky brown, darker near costal margins, veins and stigma black. Mctasoma or- C. T. 1898. On some new of Indian ange, bases of Tl to T5 more \ellowish, fading to deeper orange Bingham, species Hvmenoptera. of the Bombay Natural HistoiT Society 12: 1 l;i-130, in ]5osterior zones. Journal pi. A. i lolotvpc female: INDIA: Kainataka State: Appangala, Januan to June, 1978 (O. P. Dubev). We are indebted to Dr. B. Mallik of Cameron, P. 1902. Descriptions of new genera and species of Hv- the L'nixersity of Agricultural Sciences, Bangaloi e, India, for mak- menoptera collected bv Major C. G. Nurse at Deesa, Simla and available the of T. it is at his re- ing specimen simplex: de]30siled Ferozepore, Part If |(iuiii.il of the Bombav Natural History So- quest in the National Pusa Collection, Division of Entomolog\', ciety 14: 419-449. Indian Agricultural Research Institute, New Delhi, f^r. Mallik has Cameron, P. 1905. On the Hvmenoptera of the Albany Museum, provided fui tliei details about the collecting site. The specimen Grahamstown, South Africa. Records of the Albany Museum 1: was actuallv collected on 1 June 1978. Appangala is a small village 185-265. about 40 km from Mercara in the Coorg district of K;irnalaka, T. D. A. 1904a. etc. Emo- 75°4()'22"E, 12°32'35"N, altitude about 1 100 m; rainfall about 2:')00 Cockerell, The genus Apisla. Canadian mm amiually; vegetation wet deciduous. mologist 36: 357. 326 The Uni\'ersity of Kansas Scienc ;e Bulletin

Cockeiell. T. D. A. 1904b. The bees of soulhcni California.— 1. Mavromoustakis, G. A. 1958. Bees (Hymenoptcia,.\p(jidea) of At- Bulltiin ol llie Southern California Academy of Sciences 3: 3- tica (Greece), Part I. Ainials and Magazine of Natural Histon- 6. (13)1:433-447. Cockerell, T. D. A. 1930. Mrican bees of the family Megachilidae Ma\i omoustiikis, d. \. 1963a. On tlie bees (H\inenoplera, Apoidea) (Anthidiinae, Sempisia Aud l.ithurgus) mainly from Liberia and of Lebanon, Part III. Annals and Magazine of Natural Histoiy the Belgian Congo. Revue de Zoologie ct de Botaniqne Africaines (13)5: ()47-655. 19: 43-55. Mavromoustakis, G. A. 1963b. Further contributions to our knowl-

Cockerell, T. D. A. 1931 . Heriadine and related bees from Liberia edge of the Ethiopian Anthidiinae (Hymenoptera: Apoidea) and and the Belgian Congo. Revue de Zoologie ct de Botanique their classification.—Part 2. Annals and Magazine of Natural His- ,\fricaines20:331-34L toiy (13)6: 481-499.

Cockerel!, T. D. A., and W. P. Cockei ell. 1 901 . Contributions from Michener, C. D. 1948. The generic classification of the anthidiine the New Mexico Biological Station. — IX. On certain genera of bees. American Mu.seum Novitates 1381: 1-29. bees. Annals and Magazine of Natural Histoid (7)7: 4(>50. Michener, C. D. In press. Some genus-group names of bees. Jom- iii-xiv + + Fabricius, J. C. 1804. Systema Piezatorum. 15-349 1-30 nal of the Kansas Entomological Society. pp. Braunschewig. Morawitz, F. 1875. Mellifera Latr., pp. 1-160. In A. P. Fedtschenko, in Fairmaire, L. 1858. Ordre Hymenopteres, pp. 263-267, figures on Reisen Turkestan, Zoogeographische Folge, Teil V, pt. I, Ap- pi. 10, in]. Thomson, Voyage au Gabon, Histoire Naturelle des idae Genuinae. Moscow.

Insectes et des Archives vol. 2. 1961 . zur Kenntnis der Arten Arachnides, Entomologiques, Noskiewicz, J. Beitriige palaarktischen Friese, H. 1895. Die Bienen Europa's, Theil 1, pp. 1-128. Berlin: der Gattung Sldis. Polskie Pismo Entomologiczne 31: 1 13-133. Friedlander. Panzer, G. W. F. 1804. Faunae Insectorum Germanicae. Heft 86.

1 F. Friese, H. 1898. Die Bienen Euiopa's, Theil W, pp. 1-303, pi. Inns- Panzer, G. W. 1806. Kiitische Revision der Insektenfauna bruck: C. Lampe. Deutschlands, [14] -1-271 pp., 2 pis. Niirnberg. Friese, H. 1902. Zwei neue Bienengattungen (Hym.). Zeitschrift Pasteels, J. J. 1968. Statut, ivffinites et origines des Anthidiinae fiir Systematische Hvmenopterologie unci Dipterologie 2: 186- paiasites. Naturaliste Canadien 95: 1055-1063. 187. Pasteels, J. J. 1969a. La systematique generique el subgenerique Friese, H. 1905. Die WoUbienen Afrikas. Genus Anlhidium. des Anthidiinae (Hymenoptera, Apoidea, Megachilidae) de Zeitschrift fiir Systematische Hvmenopterologie und Dipterolo- I'ancien monde. Memoires de la Societe Royale d'Entomolo- gie 5: 65-75. giede Belgique31: 1-148. H. 1908. 5. 119-168. In Y. Wi.s- 1969b. Anthidiinae Friese, Apidae, pp. Sjostedt, Pasteels, J. f. New (Hymenoptera, Apoidea, senschaftliche Ergebnisse der schwedischen zoologischen Ex- Megachilidae) from the Mediterranean area and from the Near pedition nach dem Kilimanjaro, dem Meru und den East. Israel Journal of Entomology 4: 409-434. Deutsch-Ostafrikas 19()5-190(i, 1972. Revision des \iithidiinae umgebenden Massaisteppen Pasteels, J. ]. (Hymenoptera Band 2, Abt. 8 (Hyment)ptera), Pi. 5 (Apidae). Stockholm: Apoidea) de la region indo-malaise. Bulletin et Annales de la PalmquisL Societe Royale Beige d'Entomologie 108: 72-128. Bul- Friese, H. 1911. Apidae L Megachilinae. DasTierreich, Lieferung Pasteels, J. J. 1980. Revision du genre £i/a.5^w Gerstaecker. 28: xxvi + 440 pp. Berlin: Friedlander. letin et Annales de la Societe Ro\'alc Beige crEntomologie 1 16: Friese, H. 1923. Die europiiischen Bienen. Beilin and Leipzig: 73-89. xvii + Gruyter. 456 pp., 33 pi. Pasteels, J.J. 1984. Revision des Anthidiinae (Ihnienoptera, Gerstaecker, A. 1857. [Bees and wasps collected in Mozambique]. Apoidea, Megachilidae) de I'Afrique subsaharienne. Memoires Monatsberichte, Akademie der Wissenscliaften. Berlin, xom de la Classe des Sciences. Academic Royale de Belgique (2) 19(1): 29, October 1857, pp. 460-464. 1-165. Gislel, J. N. F. X. 1848. Naturgeschichte des Thierreichs, fiir Popov, V. B. 1941. Notes on DknUliulium \ilini(uni (Eversm.) and + Hohere Schulen. xvi 216 pp., 32 pis. Stuttgart: Hoffmann. a new species of Stdis Panz. Entomologisk Tidskrift 62: 222-224. Isensee, R. 1927. A study of the male genitalia of certain anthidi- Popov, V. B. 1950. Generic groupings of the mid-Asian bee sub- 70: inc bees. Annals of the Carnegie Museum 17: 371-382, pi. xxxii. family Anthidiinae. Doklady Akademii Nauk SSSR 31.5-318 Klug, F. 1807. Vergleichung der Galtungen der Hautfliigler, [in Russian]. Piezata Fabr. Hvmenoptera Linn. |ur. Magazin fur Insekten- Popov, V. B. 1956. New and little-known bees from central Asia. kunde 6: 189-198. Entomologicheskoe Obozrenie 35: 159-171 [in Russian].

Krombein, K. V. 1951 . Additional notes on the bees of the Solomon Radoszkowski, O. 1874. Supplement indispensable a I'arlicle pub- Islands. Proceedings of the Hawaiian Entomological St)cietv lie par W. Gerstaecker in 1869, sur quelc|ues gemes d'H\inenop)- 14: 277-295. teres. Bulletin de la Societe Imperiale des Naturalistes de Moscou Lepeletierde Saint-Fargeau, A. L. M. 1841. Histoire Naturelle des 48(1): 132-164. Insectes—Hymenopteres, vol. 2, pp. 1-680. Ritsema, C. 1874. [Zodion, Hvmenoptera, etc.].Tijd.schrift\'oor En- Mavromoustakis, G. A. 1939. New and little-known bees of the sub- tomologie 17: lxviii4xxv.

family Anthidiinae (Apoidea) . —Part I. Annals and Magazine of Roig-Alsina, A., and C. D. Micliener. 1993. Studies of the phylogeny Natural History (11)3: 88-97. and cla.ssification of long-tonguefl bees. L'niversitv of Kansas Sci- Mavromoustakis, G. A. 1951. Further contributions to our knowl- ence Bulletin 55: 12-1-162. edge of the Ethiopian Anthidiinae (Hyinenoptera: Apoidea) and Roinankova, T. G. 1988. A new far-eastern bee of the tribe An- their classification. Annals and Magazine of Natinal HisloiT thidiini. Vestnik Zoologii 1988(4): 25-30. (12)4:962-981. Smith. F. 1854. Catalogue of the H\inenopterous Insects in the MavTomoustakis, G. A. 1953. New and little-known bees of the sub- Collection of the British Museum, Part 2: 198-465, pis. \ii-xii. family Anthidiinae (Apoidea).—Part VI. .\mials and Magazine Spinola, M. 1808. Insectorum Liguriae Species Novae aul Rariores. of Natural Histoiy (12)6: 834-840. Vol. 2, ii + 262 pp. Genuae: Giavier. Old WoRi.n Axtiudiim: Bees 327

Timl)erlakc. P. H. 1941. Ten new species of .SV^'fc from California. Warncke, K, 1980. Die Biencngattung Aiilh/ilnnii Fabriciiis, 1(S04 Journal of tlie New ^brk Entomological Society 49: 123-137. in der Westpalaarktis unci in tiirkeslanisclien Becken. Ento- Tkalcu, B. 19(i7. Bemerkungcn /in Taxonomie einiger palaark- molaiina 1: 1 19-209. tischer Arten der Familie Megachilidae. Acta Entomologica Bo- Warncke, K. 1992. Die westpalaarktischen Aiten dcr Bienengat- hemoslovaca 64: 91-104. tung .S'/c/h Panzer, 1806. Enlomofauna 13: 341-376. Urban. D. 1992. Gnalhaitlhiilium, gen. n. de Aiithidiinae da America do Sul. Revista Biasileira de Zoologia 9: 337-343.