ISSN 0968-0454 | Bulletin of _ The Natural History Museum | Pee 18 JUN 1998 | : PRESENTED } GENERAL LIBRARY | | f : ( | A) \ aed | THE ' NATURAL | HISTORY i MUSEUM ! | | fi yf hi} The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History) ), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Entomology Series is produced under the editorship of the Keeper of Entomology: R.I. Vane-Wright Editor of Bulletin: Dr Gaden S. Robinson Papers in the Bulletin are primarily the results of research carried out on the unique and ever-growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. 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(Ent.) © The Natural History Museum, 1998 Entomology Series ISSN 0968-0454 Vol. 67, No. 1, pp. 1-152 The Natural History Museum Cromwell Road London SW7 5BD Issued 25 June 1998 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Bull. nat. Hist. Mus. Lond. (Ent.) 67(1): 1-64 Issued 25 June 1998 Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus Silvestri associated with the ant genus Acropyga Roger and a review of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococcidae) THE NATURAL HISTORY MUSEUM 18 JUN 1998 D.J. WILLIAMS__ GENERAL LIBRARY Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 SBD. CONTENTS PATIO GICH OM) sxe ceeussscaseivdsceasvene caanseasissesciaiassnsscesosdeveteegecsdetepar doneorae cack coSeResene aaayeGs cane teensaesecesees 2 Mite nral AM GITMC lM OGS: 1. .tac. vs cncancus-taengascanssancis+sas \ \ a AN Ni ita a WY : i ve IN \ vit ‘i AN NW Wy a \\\ aT AN nN \\ A S\\\ ‘ \ ANN \\\\ AN YS \ A NN AAN WY: WAN “i AY WN ANY Nit AN aN \\ \ WN \\ AN NAS in LN AWN is NY \\\ AN WANG iN AN AA HAN WN IAN KAY AN NOK ANN Ay) AY \ \\ AN va Y \\\y \\y' \’ Wy wy \ \\\ S\N \ Wh Nh yh Ww AAS aN NW WHIRL NS \ WK A \i \\ AY Sh AY Wi nh! ‘ LA AAW \ ANY ANA Wh \ WHA AN WW YY VAAN WN ‘ NN AY Nl AY ayy \\ \ AAAS DANY \ WN Ny \ Wath Wea A ANY NY WY Wt \ NORA) \\ \ \ vA SU A Wh 4 \ YS We ANAS | 4 wt ih ln -[- ‘ WW\ \ \\ iy! Til Vi I (| ' Ai \ en ! wy aN ARR ANE NS HWW \ hei HIN f / areas AN Ly Cet ily val ma Lod it I\Y AC NA 4 “ i uF me will iN) Cee, ae VEN qf li} ~ AE IT A Liaise eo it V, en Ny \\ \ \ aN Rites HVA ze UMN tile. iy, e eae oa os ce a Vax ee RRO a IW Fig. 6 Eumyrmococcus kusiacus sp. n. Adult female. 40 is ye cietc bts tif, GEG I LENG ohn Leathe Wh ih j CAAA 7 AAA > Lebo! Uh IW NMG Gls VAAL A LR HL fp eh NLU 1 G1 OGL iY pl l4g WG) \ \ Fig. 7 Eumyrmococcus lamondicus sp. n. Adult female. Y, S VAN A QQ) NY “ D.J. WILLIAMS MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Pp i Pac AS} k len Sr i f Goce x7 ile oF Ad My re a t Peph ep ft Li Rat a) . ATR ne aM aa NG inte ji ra aviay negeart U Dl Aare) AW aire as eee ho MM Ece RAAT Pe a i pee i es ) w/'7 VU NS Pj ) - N Vii Lhe Ris rip yy. thitlyrs\e ant afcie I Pay wish al | ahr pile Lie i TA AAA ee Hh Hi Linas TY fe if ae rlipcli Vy | Lia, ; ZZ a ra lak y DJ.W. Fig. 8 Eumyrmococcus lanuginosus sp. n. Adult female. 4] 42 D.J. WILLIAMS Fig.9 Eumyrmococcus maninjauensis sp. n. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI 4 fam , 1 Pe UA) 1 ee ANN ater PLT (hays if witen aaa AT bi v Han nf anh Hh Na \ i Ha) Mf MA Ht sath ETAL ASR RON itt NAN | AAT aye 1) Lie or reat te ir! NEN ars me A BEA AHA a AO nif ih ANON CAAA q ‘ ‘ ul Mat i RUNS NORA TORE ie A SO UR Lettnpientess it AN set ek ait Wi vi ne i ie Kp fapbuetp ey wt \s tit f NUN eh Get yi CAPITA FANT { Pa ON aA ENR Hy CAR wit we rat fi ON) " yynieyd , % ante HARA MAY RH ROR ity i AAAS ARA ARR AAAS SAMA anette OT fay Mn iN ayaa at ert nhl Wt nti} H mit a why Ra a, irae UMM pity | WM Aah gar nyt hwy ‘anna ey I Pr IATL URAL ATA RTT R ORS DOR aaa re / fiji anny Wait ly cota ALTON ea , ii Lye \ na Ni ut i f . AN Art Me ry nt 1 a a hth PRATT Mey ty Hy wn na \ uM PMA Nye @ DANA iN iy Voy i wi ant NN y ia hy hy { Fit te me Fig. 10 =Eumyrmococcus neoguineensis sp. n. Adult female. 44 D.J. WILLIAMS Zi: i heey Pt NA na a Ki ONG \ CaN TaN Baw iP aN shew 7 7 / a 4 7 7 Z ~ “= iw f \ - ~N “N Pa _ _ i ‘ XN N x . ~ ‘ D.J.W. Fig. 11 9 Eumyrmococcus nipponensis Terayama. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI (fitter y J \ MRAP LAG) t Fe ALOT oh MTA ata (Vinny NN aad TARE ANH Rua i It yl 1) i V | Li} HA EUAN EE | Filip Hertha ye ahve rye WV wernt Nu HANS fl has ee = \\ HTN j AANA | Ni wh AN F | a zg = ee eS TAN es itt Fig. 12} Eumyrmococcus queenslandicus sp. n. Adult female. 45 46 D.J. WILLIAMS C aS S SS . X a= — —— ~ = y inl mY ~ =a gL! Ui ~ i NTA + ih ~ ee iF /. ne ey x SS SSS 4 = ~ to WANG ; H = Vid )\ \ mgt th FY j u uy = Mf 1 Vi ‘ we ATI NN NH RA \ hat Front h AT Ml ee Mr tt Loe vir! wit! Tite Viti p gly, Nye iy ee USS = — rili' ~ < Ne i Nea ia EPA VEG a Mal hs S \ ai N Si atl tatiana Titi ~ Nau ~ S X = S ~ ~ TAWA f ANH RN Hifi WVtPSR 7 { i] Vr a _ Ss C - if ~ A Till ht Ss Sr Lily (yin ~ ~ Hy = PAI a & x Gre ~ a ; 3 Fig. 13. Eumyrmococcus recalvus sp. n. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Fig. 14 Eumyrmococcus sarawakensis sp. n. Adult female. D.J. WILLIAMS 48 Fig. 15 Eumyrmococcus sarawakensis sp. n. Adult male. 49 MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI CE, a Z / VY ( ‘ \ - i ( a\j i ‘ al = i —o— ——_ ¢ iS : Jc L eo | =e | Sage ao ee '. NS ee Sse \ mee eee me < ae | OE SSSe= AS \ ey ‘ Fig. 16 Eumyrmococcus scorpioides (De Lotto). Adult female. D.J. WILLIAMS 7: AH AM Mil mit at eS ks . AY NS NN & \\ me IAS AA \\ Was a a A ‘\ S INT NAS \ AK hex \ i\ enn i i NY AW sua NNER TE lh ed i uh Aan ttt) i inna al Aiea gar A Ty A SN etintatiny i / S CA Us Fig. 17 Eumyrmococcus smithii Silvestri. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Ft) y bgt A ian] if f j yy eee aa a aa } HEN ‘ li aes wy Pea Fig. 18 Eumyrmococcus sulawesicus sp. n. Adult female. all 52 D.J. WILLIAMS it Le Zz Vikt E WV = - 2S = 3 S— wag a ee i [Sees 2 s y j & LD Vy 5, ri i‘ ae yh ut - | ’ 2 zZ Ze S S = x < < ~ ~ “N S < < < < x ~ S mS = Fig. 19 Eumyrmococcus taylori sp. n. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI D.J.W. Fig. 20 Eumyrmococcus taylori sp. n. Second instar. 1 7 4 ¢ pe a/ W a Va 53 54 Fig. 21 Eumyrmococcus taylori sp. n. A. Female pupa. B. Male prepupa. C. Male pupa. D.J. WILLIAMS MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Fig. 22 Eumyrmococcus taylori sp. n. Adult male. 56 Wt carat Leg apy eens A iph fit PAP TT Fig. 23 Xenococcus acropygae sp. n. Adult female. D.J. WILLIAMS MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI D.J.W. de sp. n. First instar. 58 Fig. 25 Xenococcus acropygae sp. n. A. Female pupa. B. Male prepupa. C. Male pupa. D.J. WILLIAMS MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Fig. 26 Xenococcus acropygae sp. n. Adult male. 59 60 D.J. WILLIAMS Ut PRiy ul anit allt TU me 7 Tyg Cl mnt TULA Ve A Fits alr TMT ia Hiysthy 1 aut i K TANS it a Aa i a ARAN) GIN ea aa TTL THY AA ALTA eH TTA NY, f Fig. 27 Xenococcus annandalei Silvestri. Adult female. MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI Fig. 28 Leptorhizoecus deharvengi sp. n. Adult female. 61 D.J. WILLIAMS ‘snod0I0ULAKUN JO satdads jo dew uonNgINsIq 67 “SIA Mojany “LI SNIISAMDINS “7 O] HYNUs “FT CI saploidioos “7 p| SISUBYDMDADS “FT ¢] snajp2a4 “J TZ] snoipupjsuaanb “FZ || sisuauoddiu “7 (| SIsuaaUInsoau “FT 6 sisuanviuiupu “J g snsourgnup] “J 1, snoipuoun] ‘yg SnovISny “y ¢ sIsuaIndy “7 p apaDsunquojoy “FT ¢ snsojnoi20f “q 7 SNIDIYIUIAOD “7 | NS ON S 63 MEALYBUGS OF THE GENERA EUMYRMOCOCCUS SILVESTRI AND XENOCOCCUS SILVESTRI ‘sno2020uaX JO saidads Jo deul uonngmnsIq = O¢ “SIA lajppupuun “¥ 6] av3sddo.ov “¥ 3] D.J. WILLIAMS INDEX Synonyms and misidentifications are in italics; main citations in bold acropygae 4, 23, 31 albidus 4, 5 andensis 29 annandalei 23, 25, 31 annandalei 23 atopoporus 28 Brevicoccus 26, 30 caladii 4, 31 caldasiae 4, 27, 31 Capitisetella 26, 30 Chavesia 27 clavisetosus 26 cobelopus 29 Coccidella 29 cocois 30 coffeae Geococcus 4, 31 coffeae Rhizoecus 4, 29, 31 colombiensis 29 corinthiacus 9, 10, 30 deharvengi 27 Eumyrmococcus 8, 30, 31 eversi 4, 5, 31 falciculosus 9, 10, 30 falcifer 4,5, 28, 31 Geococcus 26, 30 globosus 29 hawaiiensis 29 incrassatus 29 kelloggi 29 kolombangarae 9, 11, 30 kruiensis 9, 11, 31 kusiacus 9, 12, 31 lamondicus 9, 12, 31 lanuginosus 9, 13, 31 Leptorhizoecus 27, 30 maninjauensis 9, 14, 31 migrans 4, 26, 31 Morrisonella 29 morrisoni 28 moruliferus 4, 31 Neochavesia 27, 30 neoguineensis 9, 14, 31 Neorhizoecus 29 nipponensis 9, 15, 31 Pararhizoecus 28 petiti 28 poensis 29 poltavae 29 Prorhizoecina 26 Prorhizoecus 28 proximus 4, 28, 31 Pseudorhizoecina 26 Pseudorhizoecus 28, 30 Pygmaeococcus 28, 30 queenslandicus 9, 16 Radicoccus 29 radicum 26 recalvus 10, 17, 31 Rhizoecina 26 Rhizoecinae 2, 26 Rhizoecini 2 Rhizoecus 28, 30 Ripersiella 28 rumicis 28 sarawakensis 9, 18, 31 scorpioides 9, 19, 31 smithii 9, 19, 31 sulawesicus 9, 20, 31 taylori 10, 20, 31 trinidadensis 4, 31 weber 31 Xenococcini 2 Xenococcus 22, 30 Bull. nat. Hist. Mus. Lond. (Ent.) 67(1): 65-78 Issued 25 June 1998 Monophyly of the dacetonine tribe- group and its component tribes (Hymenoptera: Formicidae) BARRY BOLTON Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 SBD, UK (e-mail: bb@nhm.ac.uk) CONTENTS MRO CLIN C HOM copes sso en snaps os pacndids3.0. The reason why these three non-dacetonine taxa have a similar MTI range to the dacetonine-group MONOPHYLY OF THE DACETONINE TRIBE-GROUP is that whilst their heads have remained plesiomorphically broad anteriorly, their toruli have secondarily migrated outwards on the cephalic dor- sum, bringing them into line with the clypeal intersection points of the mandibles. Thus two com- pletely different evolutionary routes, static toruli combined with narrowed anterior head (the dacetonine- group), versus static broad anterior head combined with outwardly migrated toruli (Tatuidris, Cataulacus, some Cephalotini), will give the same overall MTI result. For an illustration of this character spanning the entire subfamily compare the full-face view photo- graphs of the various myrmicine genera in Bolton (1994). As a direct result of the narrowing of the anterior head the dacetonine-group shows a number of other derived characters that are unavoidable corollaries of this modification. 1. Width of labrum at base, and consequently also of clypeo-labral hinge, is reduced (see below). 2. Width of buccal cavity, and consequently also of labio-maxillary complex, is reduced (compare Fig. 3 with Figs. 12, 26, 28). 3. Anterolateral surface of head tends to be vertical, or nearly so, and is very close to or more or less directly below the antennal socket (e.g. Figs. 11, 182 19532): Preocular carina is present With the head in profile or dorsolateral view there is a ridge, carina or lamella that originates at the posterola- teral termination of the clypeus, extends posteriorly below the antennal socket and usually continues for some distance towards the level of the eye (Figs. 11, 18, 19, 30, 32). In many taxa this preocular carina is also visible in full-face view, at least anteriorly (Figs. 13, 23, 25, 31), but usually cannot be seen in this view in those species or genera where the frontal lobes and frontal carinae are strongly expanded laterally (Fig. 7). Morphologically the carina represents the exagger- ated outer margin of the antennal fossa and is another corollary of the narrowing of the head (particularly of number 3, immediately above). At its weakest the carina 1s a ridge that emphasises the dorsalmost line of the more or less vertical side of the head capsule. At its strongest it is a broad, laterally projecting lamella. The structure is perhaps better termed the subtorular carina or subantennal carina as this area is always where it is strongest developed, but earlier literature has used preocular carina so much that the name will probably be retained. Specialised range-finder/trigger hairs are developed on the mouthparts In the dacetonine-group hunting always seems to in- volve an open-mandible approach to the prey, followed 69 by a rapid strike with the mandibles (see references above, particularly Masuko, 1985). The strike may be followed by static pressure of the mandibles, merely to retain a grip on the prey until the sting can be brought into use (‘strike-hold-sting’ technique), or dissipation of the kinetic energy of the strike itself may be suffi- cient to shock the prey into immobility, even if only temporary, until the sting can be brought into play, particularly if the prey is lifted clear of the ground immediately after the strike (‘strike-lift-sting’ tech- nique). Whatever the technique, the explosive closing of the mandibles is initiated by activation of a trigger which consists of one or more specialised hairs on the mouthparts. Contact with prey by these hairs com- mences a sequence of events that causes the mandibles to snap shut (Masuko, 1985; Gronenberg, 1996). In the vast majority of dacetonine-group taxa these hairs arise from the anterior margin of the labrum or the apices of the labral lobes (Figs. 16, 25-31 (broken off short in 26), 27, 33), more rarely from the mandible itself; such trigger hairs are not found elsewhere in the Myrmicinae. Specialisation of the labrum Except for the dacetonine-group of tribes the structure of the labrum is very uniform and generalised in the Myrmicinae (Gotwald, 1969 and present investiga- tion), and this same structure is prevelent throughout the Formicidae. Outside the dacetonine-group the la- brum is a simple sclerite (Fig. 3) that is broader than long and broadly hinged to the clypeus. Its posterior margin, which is attached to the clypeus, is more or less straight. Its anterior (free) margin is indented or cleft medially so that in general the labrum appears broadly B-shaped, or D-shaped with a median inden- tation in the anterior free margin. The clypeo-labral hinge is mobile so that at rest the labrum folds back and down with respect to the clypeus. In this position it tightly overlaps and protects the delicate apical portions of the labio-maxillary complex. In the Myrmicinae this is the plesiomorphic shape and posi- tion. It is encountered in all the morphologically more generalised tribes (e.g. Myrmicini, Tetramoriini, Pheidolini, Pheidologetonini, Formicoxenini, Solenopsidini), as well as in those that are rather more specialised morphologically (e.g. Attini, Cataulacini, Cephalotini, Crematogastrini, Stenammini, Agroecomyrmecini). Tribes in the dacetonine-group show a number of modifications away from the generalised structure illustrated in Fig. 3, but all are based on labral narrow- ing and elongation, and loss of the basic broadly B- or D-shaped outline. In taxa with mandibles that use static pressure, regardless of whether the mandibles are long or short, the labrum becomes longer and narrower, sometimes longer than broad. The sclerite usually terminates in 70 one or two exaggerated anterior lobes or linguiform prominences that bear the trigger hairs (Figs. 12, 16, 25, 28, 29, 31). In these forms the labrum can usually still flex down very slightly, but it is not capable of tight closure against the labio-maxillary complex (Figs. 12, 28). Conversely, in some taxa the labrum is hyper- trophied and elongate-linguiform; instead of concealing only the apex of the labio-maxillary complex it covers the entire buccal cavity (Fig. 32). Taxa with long kinetic mandibles tend to have the labrum very narrow basally, and more distally to develop a pair of laterally projecting processes or arms; the sclerite therefore tends to be roughly T-shaped and its apex functions to prop open the mandibles prior to striking (Figs. 26, 27, 30). One long-mandibulate genus (Acanthognathus) has eliminated the labrum from this function and here it is represented only by a very slender, inverted Y- shaped sclerite. Some characters considered but not used at tribe-group rank Presence of 2-segmented antennal club An antennal club of two segments is universal in the tribes Basicerotini, Dacetonini and Phala- cromyrmecini, though only very poorly expressed in Daceton. This may be the result of a single evolution- ary event in the dacetonine-group, but a two-segmented club is also developed in a wide range of non- dacetonine myrmicine taxa. For example, a strongly defined two-segmented club is universal in Melissotarsini (Melissotarsus, Rhopalomastix), present in all the core genera of Pheidologetonini (Afroxyidris, Carebara, Oligomyrmex, Paedalgus, Pheidologeton), present in some but by no means all Stenammini (Adelomyrmex, Baracidris, Lachnomyrmex, Mayriella, Tetheamyrma), occurs in a few Solenopsidini (Carebarella, Solenopsis) and is present in the single extant genus of Agroecomyrmecini (Zatuidris). It is weakly developed in some Blepharidattini (Blepharidatta, Wasmannia), and in some but not all species of Cardiocondyla (Formicoxenini) and Crematogaster (Crematogastrin1). Antennomere count Although very useful as acharacterin keys, antennomere count was ignored in this survey because in some myrmicines there is variation within genera (e.g. 10, 11 or 12 in Tetramorium (Tetramoriini); 4, 5 or 6 in Smithistruma (Dacetonini)). Also, within the tribes of the dacetonine-group there is a wide range of counts that makes analysis very difficult. For example in Dacetonini counts of 4, 5, 6 and 11 have been recorded; in Basicerotini 7,8,9 and 12, andin Phalacromyrmecini 8, 9 and 11. Iam convinced that these counts indicate independent morphoclinal reductions within each tribe, with no significance at tribe rank. B. BOLTON Absence of tibial spurs Throughout the tribes of the dacetonine-group spurs are absent from the middle and hind tibiae. This condi- tion is apomorphic among the Myrmicinae but unfortunately is so extremely widespread through the subfamily that it has very little analytical value in the current survey. MONOPHYLY OF TRIBE BASICEROTINI Basicerotini Brown Basicerotini Brown, 1949c: 86. Type-genus: Basiceros Schulz, 1906: 156. [Basicerotini relegated as junior synonym of Dacetonini by Baroni Urbani & de Andrade, 1994: 10; revived from synonymy by Bolton, 1995b: 9.] Apomorphies of tribe Basicerotini Each worker apomorphy is briefly described, in some cases with added comments where unrelated taxa show similar but non-homologous modifications. The plesiomorphic state of each character is given in square brackets; in some cases alternative apomorphic condi- tions are also noted for comparative purposes. LABRUM Distal of the hinge with the clypeus the labrum has a deeply incised transverse groove or trench across its entire dorsal width. The distal margin of this groove is defined by a sharp edge, ridge or crest (Fig. 16; also visible between the closed mandibles in Figs. 13, 14). {Labrum without sharply defined transverse groove distal of hinge. ] In the basicerotines this groove, or at least its sharp distal margin, is usually located far enough anteriorly on the labrum as to be visible in ordinarily mounted specimens which have the mandibles ajar and the labrum slightly depressed. TORULUS The dorsal lobe of the torulus is hypertrophied and strongly curved downwards (Figs. 18, 19); its outer surface is nearly vertical so that it conceals part to most of the condylar bulb of the scape and the antennal socket itself. [Torulus a simple annulus or with a small lobe present dorsally that is horizontal and does not conceal the condylar bulb of the scape and the antennal socket. | SCAPE NECK ARTICULATION Because of the size and shape of the torulus the space in which the basal neck of the scape (the short narrow section between condylar bulb and scape shaft proper) MONOPHYLY OF THE DACETONINE TRIBE-GROUP can move is narrow and directed fore and aft (Figs. 18, 19). [Space in which basal neck of scape can move is not restricted to a narrow fore and aft motion. ] BASE OF SCAPE Scape shaft near base is bent downwards through a right-angle or near right-angle; the articulatory condyle at the extreme base projects forward from this through another right-angle that is not in the same plane as the first bend but rather is rotated through about 90 degrees (Figs. 8, 9). [Scape shaft in a straight line with basal condyle; or if scape angled downward near base then angle of basal condyle remains in the same plane as the shaft and is not rotated through 90 degrees. ] ANTENNAL FOSSA Antennal fossa separated from scrobe by at least a cuticular rim or crest; the depressed fossal area sur- rounded on all sides by raised or prominent cuticle (Figs. 18, 19). [Antennal fossa and scrobe confluent (when the latter is developed). ] SCROBE Scrobe always present and located below the eye (Figs. 18, 19), the latter usually situated on the extreme dorsolateral rim of the scrobe, more rarely towards underside of upper scrobe rim (very rarely eye absent). [Scrobe absent, or present but extending above the eye (eye sometimes absent). ] Some other myrmicine taxa have a scrobe that extends below the eye, namely Cataulacus, some Cephalotini, and the Dacetonini related toEpopostruma (Epopostrumiti in the synoptic classification above). Cataulacus and the few cephalotines with this condi- tion can be dismissed as obvious convergence. Not only is the basic structure of the scrobe dissimilar but also the detailed structure of the head is very different. They lack, of course, the apomorphies of the dacetonine-group of tribes and exhibit their own series of apomorphies. The Epopostrumiti, on the other hand, belong in the dacetonine-group. Apart from possess- ing the apomorphies of Dacetonini and lacking those of Basicerotini, the scrobes in Epopostrumiti lack sharply defined posterior margins, usually lack vent- rolateral margins and, except in a very few species, fail to extend forward to the mandibular articulation (Fig. 32); all of these are developed in Basicerotini. Finally, a morphocline of species in the genera Colobostruma — Mesostruma — Epopostruma is present that exhibits a gradual increase in development and definition of the scrobe; no basicerotine could be inserted into the series, nor added to either end. OCCIPITAL FORAMEN The occipital foramen is set in a deep depression on the occipital surface of the head; cuticular margination is continuous around the depressed area (Fig. 17). [Oc- cipital foramen not set in a continuously marginate deep depression. ] Fl A very few Dacetonini have a transverse ventral rim of cuticle below the occipital foramen. This structure is very different from the basicerotine organisation. HELCIUM Helcium arises from the base of a broad, deeply con- cave depression or excavation in the anterior surface of the postpetiole (Fig. 20). [Helcium not set in a concave depression. | GASTER First gastral tergite and sternite each distinctly trans- versely marginate basally, immediately behind the postpetiole (Fig. 21). [First gastral tergite and sternite not marginate basally. ] Members of the strumigenyite group of Dacetonini genera have a specialised transverse crest on the first gastral tergite, the limbus, which is an apomorphy of that group. The limbus is located prebasally and is inclined towards the base proper, where it overhangs the presclerites of the segment. Itis nota homologue of the basal margination developed in Basicerotini. No Dacetoninihavea basally marginate first gastral sternite. SCULPTURE First gastral tergite and sternite with characteristic sculpture of dense, sharply incised, separated punc- tures (e.g. Fig. 21). [First gastral tergite and sternite unsculptured, or with different sculpture. | This character may seem rather imprecise, but the form of sculpture is striking and not obviously re- peated elsewhere. A very few individual basicerotine species have secondarily reduced or effaced the sculp- ture. Elsewhere in the dacetonine-group, and in the Myrmicinae as a whole, are many species with sculp- tured gasters. Even when the sculpture in these is of a basically punctate form it tends to be reticulate-punc- tate or sparse, or on one sclerite but not the other. It is not the dense deeply-incised punctation exhibited by the Basicerotini on both sclerites. MONOPHYLY OF TRIBE DACETONINI Dacetonini Forel Dacetonini Forel, 1892: 344. Type-genus: Daceton Perty, 1833: 136. Apomorphies of tribe Dacetonini The two worker apomorphies are briefly described and comments added. The plesiomorphic state of each character is given in square brackets. MANDIBLE Mandible with a medially projecting cuticular process present on the inner margin close to the base 72 (basimandibular process); the process not merely a modified tooth (Figs. 25, 26, 27, 31, 33). [Mandible without a basimandibular process. ] In some dacetonine taxa the basimandibular process is visible in ventral view in ordinarily mounted speci- mens, but in most the mandibles need to be opened quite widely or the labrum depressed. The process is variably shaped in different dacetonine taxa, taking the form of a lobe, a short or long spur, or a lamella (basal lamella). In all dacetonines except Acanthognathus the basimandibular process inserts between the clypeus (above) and the labrum (below) when the mandibles are closed and is part of the jaw locking mechanism. In Acanthognathus the process is hypertrophied but passes ventral to the labrum as the latter is vestial in this genus and no longer serves in the jaw locking mechanism. A few basicerotine species in the genera Eurhopalothrix and Octostruma have the basal tooth of the mandible flattened or lengthened. This is not homologous with the dacetonine basimandibular proc- ess, which is derived from the mandible itself and not from a tooth. Basicerotine species with a modified basal tooth are exceptional and certainly best regarded as independent acquisitions. The usual condition is to have the basal tooth quite normal; presence/absence of a flattened basal tooth cannot be used diagnostically either at genus or species-group rank. LABRUM Dorsal surface of labrum with an impression or pair of impressions located medially on the labral shield, distal of the basal hinge but proximal of the labral lobes (Figs. 27, 30). [Labrum without mid-dorsal im- pression. | The labral impression receives the basimandibular processes of the mandibles when they are fully closed. The character is not repeated anywhere else in the Myrmicinae. It is secondarily lost in Acanthognathus because, as pointed out above, the labrum is secondar- ily extremely reduced in this genus. MONOPHYLY OF TRIBE PHALACROMYRMECINI Phalacromyrmecini Dlussky & Fedoseeva Phalacromyrmecini Dlussky & Fedoseeva, 1988: 80 [based on diagnosis in Bolton, 1984: 381]. Type- genus: Phalacromyrmex Kempf, 1960: 89. [Phalacromyrmecini relegated as junior synonym of Dacetonini by Baroni Urbani & de Andrade, 1994: 10; revived from synonymy by Bolton, 1995b: 9.) B. BOLTON Apomorphies of tribe Phalacromyrmecini Each worker apomorphy is briefly described and com- ments added. The plesiomorphic state of each character is given in square brackets. MANDIBLE Dentition of alternating large and small teeth from base to apex; largest tooth usually the basal. [Dentition not of alternating large and small teeth; largest tooth usually the apical. ] A few isolated species or small species-groups in the dacetonine genus Glamyromyrmex have the basal tooth the largest on the masticatory margin, but these lack alternating dentition. MESOPLEURON Katepisternum with an impression or groove extend- ing obliquely downward from posterior margin of mesopleural hair-filled gland towards metapleuron; impressed area usually bounded by ridges or carinae. [Katepisternal oblique groove absent. ] SCAPE Scape slender basally, clavate apically; entire scape roughly Indian-club shaped. [Scape subcylindrical.] POTENTIAL SYNAPOMORPHIES BETWEEN PAIRS OF COMPONENT TRIBES The object of this paper has been to establish the monophyly of the tribes and the tribe-group, rather than to produce a formal phylogeny. However, anumber synapomorphies potentially linking pairs of tribes within the group have been noticed and these are mentioned below. The plesiomorphic state expressed by the isolated tribe in each set is given in square brackets. Potential synapomorphies of Dacetonini + Phalacromyrmecini POSTPETIOLE-GASTER ARTICULATION Diameter of presclerites of abdominal segment 4 (=first gastral segment) small and constricted so that the postpetiole-gaster articulation is relatively narrow. [Basicerotini: diameter of these presclerites broad so that the postpetiole-gaster articulation is relatively wide. ] PRESCLERITE OF FOURTH ABDOMINAL SEGMENT Pretergite of abdominal segment 4 (=first gastral seg- ment) subtended by a short narrow neck-like constriction. [Basicerotini: pretergite subsessile to ses- sile.] MONOPHYLY OF THE DACETONINE TRIBE-GROUP BASIMANDIBULAR GLAND Basimandibular gland present. [Basicerotini: basimandibular gland absent. ] This gland is variously developed in many genera of these two tribes (/shakidris, Microdaceton, Strumigenys, Glamyromyrmex); it is universally ab- sent from basicerotines. Its absence from some dacetonines (Orectognathus, Acanthognathus) may be secondary. MESOPLEURAL GLAND Anterolateral angle of mesopleuron bears a hair-filled glandular structure set in an emargination of the rim of the sclerite. [Basicerotini: mesopleural gland absent. ] This supposed gland varies from absent to mas- sively hypertrophied in Dacetonini and Phalacromyrmecini. It is universally absent in Basicerotini. Potential synapomorphies of Basicerotini + Phalacromyrmecini BASE OF MANDIBLE Base of mandible with a long stiff ventrally directed seta on ventral margin. [Dacetonini: such a seta ab- sent. ] This specialised seta is present in all Basicerotini. In Phalacromyrmecini it is obvious in /shakidris. Not recorded in the single specimen of Phalacromyrmex currently available, but this is in poor condition and badly mounted. PROPODEAL SPIRACLE Propodeal spiracle low on side of sclerite, abutting the margin of the small metapleural gland bulla. {Dacetonini: propodeal spiracle high on side, widely separated from metapleural gland. | In the dacetonine genus Acanthognathus the spiracle abuts the metapleural gland bulla but here the spiracle is high on the side and the bulla is secondarily ex- tended upwards. Potential synapomorphies of Dacetonini + Basicerotini None detected. APPENDIX: Species dissected The parts of all dissected specimens, whether partially or completely disarticulated, have been remounted on green-flagged card points and are deposited in The Natural History Museum, London. [List does not in- clude the many taxa mounted with mandibles open and mouthparts displayed, but not otherwise dissected; these have been mounted on blue-flagged card points in the Natural History Museum, London, collection. ] Basicerotini Basiceros. discigera, manni, militaris, singularis. Eurhopalothrix: australis, biroi, bolaui, dubia, floridana, gravis, heliscata, insidiatrix, jennya, omnivaga, procera, punctata, speciosa, spectabilis, szentivanyl. Octostruma: balzani, betschi, iheringi, inca, rugifera, stenognatha. Protalaridris: armata. Rhopalothrix: ciliata, isthmica, plus | unidentified species. Dacetonini Acanthognathus: brevicornis, ocellatus, rudis. Codiomyrmex: thaxteri. Colobostruma: alinodis, cerornata, plus | unidenti- fied species. Daceton: armigerum. Epitritus: argiolus, hexamerus, laticeps, roomi. Epopostruma: frosti. Glamyromyrmex: beebei, excisa, flagellatus, semicomptus, sistrurus, tukultus, plus 2 unidenti- fied species. Gymnomyrmex: villiersi. Kyidris: mutica, plus | unidentified species. Mesostruma: browni, turneri. Microdaceton: exornatum, tibialis, plus | unidentified species. Neostruma: brevicornis, crassicornis, myllorhapha, zeteki. Orectognathus: antennatus, clarki, mjobergi, szentivanyi, versicolor. Pentastruma: sauteri. Quadristruma: emmae. Serrastruma: lujae, ludovici, serrula. Smithistruma: alberti, angulata, dohertyi, fridericimuelleri, microthrix, ornata, transversa, truncatidens, plus 6 unidentified species. Strumigenys: biolleyi, chyzeri, denticulata, doriae, elongata, gundlachi, godmani, koningsbergeri, louisianae, micretes, nidifex, pallestes, perplexa, precava, prospiciens, rogeri, rukha, saliens, signeae, subedentata, tigris, trieces, trudifera, plus 12 unidentified species. Trichoscapa: membranifera. Phalacromyrmecini Phalacromyrmex: fugax. Ishakidris: ascitaspis. ACKNOWLEDGEMENT My sincere thanks to Suzanne Lewis (Department of Entomology, The Natural History Museum) for taking the SEM photographs used to illustrate this paper. 74 REFERENCES Arakelian, G. R. & Dlussky, G. M. 1991. Murav’i triby Dacetini SSSR. Zoologichesky Zhurnal 70 (2): 149-152. Baroni Urbani, C. & de Andrade, M. L. 1994. First description of fossil Dacetini ants with a critical analysis of the current classifica- tion of the tribe. (Amber Collection Stuttgart: Hymenoptera, Formicidae. VI: Dacetini.) Stuttgarter Beitrdge zur Naturkunde Serie B (Geologie und Palaontologie) 198: 1-65. Baroni Urbani, C,. Bolton, B. & Ward, P. S. 1992. The internal phylogeny of ants. Systematic Entomology 17: 301-392. Belshaw, R. & Bolton, B. 1994. A survey of the leaf litter ant fauna in Ghana, West Africa. Journal of Hymenoptera Research 3: 5-16. Bolton, B. 1983. The Afrotropical dacetine ants. Bulletin of the British Museum (Natural History) (Entomology) 46: 267-416. Bolton, B. 1984. 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Myrmecological Society of Japan: 94 pp. Perty, M. 1833. Delectus animalium articulatorum Fase. 3: 125— 224. Monachii. Taylor, R. W. 1990. New Asian ants of the tribe Basicerotini, with an on-line computer interactive key to the twenty-six known Indo- Australian species. Invertebrate Taxonomy 4: 397-425. Wheeler, G. C. & Wheeler, J. 1976. Ant larvae: review and synthesis. Memoirs of the Entomological Society of Washington 7: 1-108. Wheeler, G. C. & Wheeler, J. 1985. A simplified conspectus of the Formicidae. Transactions of the American Entomological Society 111; 255-264. Wheeler, W. M. 1910.Ants: their structure, development and behavior: 663 pp. New York, Columbia University Press. Wheeler, W. M. 1922. The ants of the Belgian Congo. Bulletin of the American Museum of Natural History 45: 1-1139. Wilson, E. O. 1953. The ecology of some North American dacetine MONOPHYLY OF THE DACETONINE TRIBE-GROUP 75 ants. Annals of the Entomological Society of America 46: 479-495. Wilson, E. O. & Brown, W. L. Jr. 1985. Behavior of the cryptobiotic Wilson, E. O. 1956. Feeding behavior in the ant Rhopalothrix biroi predaceous ant Eurhopalothrix heliscata, n. sp. Insectes Sociaux Szabo. Psyche 63: 21-23. 31: 408-428. 76 B. BOLTON Figs. 1-11. Worker ants: 1-3, Head of Myrmica rubra; 1, tilted slightly back from full-face view; 2, oblique close-up of fully closed mandibles; 3, ventral view to show buccal cavity; 4, head of Cataulacus lujae, tilted back from full-face view; 5-7, heads in full-face view of 5, Pilotrochus besmerus; 6, Tatuidris tatusia; 7, Ishakidris ascitaspis; 8-9, scape of Basiceros singularis, condyle uppermost; 8, ventral view; 9, dorsal view; 10-11, head of Eurhopalothrix platisquama; 10, full-face view; 11, oblique frontal view. MONOPHYLY OF THE DACETONINE TRIBE-GROUP 77 Figs. 12-22. Worker ants: 12, ventral head to show buccal cavity of Eurhopalothrix platisquama; 13-15, heads in full-face view of 13, Rhopalothrix ciliata, 14, Protalaridris armata; 15, Basiceros discigera; 16, oblique frontal view of head of Basiceros singularis, mouthparts extended and right mandible removed; 17, occipital foramen of Eurhopalothrix heliscata; 18-19, lateral view of head with antenna removed, of 18, Basiceros singularis; 19, Eurhopalothrix platisquama; 20, frontal view of helcium of Eurhopalothrix procera; 21, base of gaster in oblique ventral view of Basiceros singularis; 22, | head in full-face view of Dysedrognathus sp. n. 78 B. BOLTON Figs. 23-33. Worker ants. 23-25, head in full-face view of 23, Smithistruma sp. n.; 24, Daceton armigerum; 25, Smithistruma reliquia, mandibles open; 26-28, ventral view of head to show buccal cavity of 26, Microdaceton sp. n., mandibles open; 27, Daceton armigerum, right mandible and labio-maxillary complex removed; 28, Smithistruma truncatidens;, 29-30, oblique frontal view of head to show labrum, mandibles removed, of 29, Smithistruma truncatidens; 30, Strumigenys sp. n.; 31, head in full-face view of Smithistruma kerasma, mandibles open; 32, head in ventrolateral view of Colobostruma sp.; 33, mandibles in oblique frontal view of Microdaceton tibialis. Bull. nat. Hist. Mus. Lond. (Ent.) 67(1):79-152 Issued 25 June 1998 An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae, Bombini) PAUL H. WILLIAMS Department of Entomology, The Natural History Museum, Cromwell Road, South Kensington, London SW7 5BD CONTENTS | Hii OTG IC COLAC 01 SA SPR ee ORE nc RRRPEY cece ce ert pe SScR PPA Rae ORME 08 dc. ORE Sm ok Ore onan eee 80 PAS EALISE SEO IES IECLES socass vaaeteraccee dr caamevusa scaversussaeroncetaxcnes cade caps sauseizeccsee ured Tatars meen gone otsers = ane 80 FUISLOUY ON CIS COVEUY OL SPECIOS fences onseteteseascsncoseesasgatusch cust daceacoasy shes eatovedadec¥oates diospsoeesucece teas 80 ANSTO yal Chin UNIT GALCL OM OL TALES) « cecps deen ce 1790), 1830) | 1870) 190 ISSO RT SS0 Fig. 2 Cumulative number of first formal descriptions of presently recognised bumble bee species with centres of area of occupancy (so species lists do not overlap) in each of the four principal biogeographic regions occupied by bumble bees (dates from the oldest available names in the sense of ICZN, 1985; regions defined in Williams, 1996b: fig. 1; Oriental includes northern and southern Oriental Regions; Nearctic includes northern, central and southern Nearctic Regions; Neotropical includes northern, central and southern Neotropical Regions; the Arctic Region is excluded; species that are exclusively peri-Tibetan Oriental but which nevertheless have range centres in Palaearctic central Tibet by simple range averaging are included as Oriental species). Pieridae combined. It must be born in mind that the insect taxa that they surveyed are all more speciose than the bumble bees by a factor of at least four, and extreme values for larger groups are less likely. None- theless, Gaston & Mound (1993) also noted that the two families of most brightly coloured butterflies have the highest levels of synonymy and that these families have many more subspecific names than the smaller and duller-coloured hesperiid butterflies. R. I. Vane- Wright (pers. com.) suggests that synonymy rates may be particularly high among the large, colourful butter- flies of the Danainae and Parnassius. In contrast to the discovery of currently recognised species, the greatest activity in publishing names for all supposed bumble bee taxa at the rank of species and below was concentrated slightly later than for pres- ently recognised species, in the first half of the twentieth century (Fig. 3, median date 1922). This difference may be explained in part by the logical inevitability that synonyms and names for taxa below the rank of species can only be published subsequently to valid species names (i.e. the oldest available names, exclud- ing junior homonyms, in the sense of ICZN, 1985). If these names were in effect to represent the redescription of known species at random, then the earlier described species might be expected to have accumulated more names. Studies of other taxa have also shown that both the date of first description and the number of syno- nyms per species may be affected by variation in the size of a species’ geographic range (as well as by other factors such as body size). Large range size is likely to affect the date of first description because it contrib- utes to a greater ‘apparency’ of the species to collectors (Gaston, Blackburn & Loder, 1995), particularly as broad correlations between range size and abundance suggest that widespread species also tend to have higher local densities (Brown, 1984; Gaston, 1994; for bumble bees, see Hanski, 1982; Williams, 1988).Apart from enhancing the chances of random redescription, large range size is also likely to affect the number of synonyms because there is a greater likelihood that specimens collected in one area will be regarded as CHECKLIST OF BUMBLE BEES 3000 83 2500 2000 Names 3 + 1000 + 1750 1770 1790 1810 1830 1850 1870 1890 1910 1930 1950 1970 1990 Date of description Fig. 3 Rate (lower grey) and cumulative number (upper white) of all descriptions with classical names for bumble bee species, subspecies and infrasubspecies since the starting point of zoological nomenclature in 1758 (from a manuscript catalogue, unpublished). distinct from specimens collected from another distant area, because character variation is apt on average to be greater (Gaston, Blackburn & Loder, 1995). For the bumble bee catalogue data, the number of synonyms (including subspecies, but excluding infrasubspecies) is correlated with both the date of first description and the range size of a species independ- ently of one another, although slightly more of the variation is explained by variation in range size (par- tial r, Table 1). Many of the species with large range sizes, early dates of first formal description and many synonyms are found in western Europe (i.e. triangles at the left and upper part of Fig. 4). Most of these species occur in either the lowland areas of Europe where early naturalists were most active, such as Britain, or else are nearly circumpolar in their distribu- tion. Curiously, all of the infrasubspecific names (34% of all names as interpreted at present) belong to the bumble bee species of the Old World (Fig. 5). Species of the Old World also have more synonyms and sub- species per species than do the species of the New World (numbers of names log-transformed and ex- cluding 6 Holarctic species, 1,,,.= 3.81 with separate variance estimates, p< 0.001). | One possible explanation for the greater numbers of names per species for bumble bees of the Old World is that they might have broader distributions than the species of the New World (see above). This could arise because the Old World has a slightly larger total area of suitable habitat (bumble bees oc- cupy 131 of the 611,000 km? grid cells in the Old World and 117 in the New), which is apparently subdivided into fewer well differentiated biogeographic assemblages of bumble bee species (e.g. Williams, 19965: fig. 1). However, this explana- tion is not strongly supported by the bumble bee data, which show the difference in range sizes between the two hemispheres to be not significant, (range sizes log-transformed and excluding 6 Holarctic species, t,,,= —1.24 with separate variance estimates, p= 0.22). Consequently, while an effect of differences in habitat area will deserve future consid- eration, other effects are likely to be more important. A second possibility is that whereas bumble bee taxa of uncertain rank may have tended to be re- garded more often as subspecies in the Old World, in the New World they may have tended to be regarded as species (see the discussion below of criteria to recognise species). While this factor could have con- tributed to the observed patterns, it is unlikely to explain why (at a lower rank) so many infrasub- specific names were described exclusively for taxa from the Old World. A third possibility is that the diversity of languages used for taxonomic publications in the Old World may 84 Table 1 P.H. WILLIAMS Results of multiple regression of numbers of synonyms/subspecific names (infrasubspecific names are excluded; from a manuscript catalogue, unpublished) on date of first formal description for presently accepted species and range size (number of occupied 611,000 km? grid cells world-wide). Partial r values indicate the correlations with the synonyms variable after adjusting for the other predictor variable in each case. log, ,(synonyms+1) = 6.316(+0.969) — 0.003(+0.0005).date + 0.401(+0.057).log, (range) multiple = 0.72 Fg leo 16 p< 0.0001 partial r Ure Dp date —0.390 -6.51 < 0.0001 log,,(range) 0.418 7.08 < 0.0001 pascuorum a lucorum s.] 4 humilis = guahuoukS soroeensjs & terrestris Fig. 4 Scatterplot of 239 presently accepted bumble bee species by range size (number of occupied 611,000 km? grid cells world-wide), date of first formal description and numbers of synonyms/subspecific names (infrasubspecific names are excluded; from a manuscript catalogue, unpublished). The British fauna is distinguished as filled triangles, the nearly circumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some British and widespread European species are labelled individually. CHECKLIST OF BUMBLE BEES 85 1200 | 1000 | 800 } 600 Numbers 400 200 Old World MMM Species Synonyms [= Infrasubspecies New World Fig.5 Number of presently recognised bumble bee species, synonyms/subspecific names and infrasubspecific names for the Old World and the New World (from a manuscript catalogue, unpublished). have impeded communication and lead to more fre- quent re-description of taxa than in the New World, where English was much more dominant (C. O’ Toole, pers. com.). Again, while this factor is likely to have contributed to the observed patterns of synonyms, it does not explain why (at a lower rank) so many infrasubspecific names were described exclusively for taxa from the Old World. Another possible interpretation, which might ex- plain more of the differences in description dates between Figs. 1-3 as well as the differences in the distribution of bumble bee subspecies, synonyms and infrasubspecific names between hemispheres (Fig. 5), is that during the twentieth century, effort for describ- ing the variety of these insects may have become, in effect, re-directed towards finer distinctions and lower nomenclatural ranks within known species. This is perhaps likely as undescribed species became inevita- bly more difficult to find close to home for the most active taxonomists, who were based in Europe. Three lines of evidence are consistent with this explanation. First, slightly more of the variation in richness of infrasubspecific names among species is accounted for by variation in the date of first description of the species (partial r, Table 2), rather than by variation in total range size. This is in contrast to the pattern for synonyms alone (cf. Table 1), although species that are sufficiently widespread in lowland Europe to include Britain within their distributions still tend to have high numbers of both synonyms and infrasubspecific names (Fig. 6, e.g. B. pascuorum, B. lucorum). A second intriguing observation is that compared to the number of authors who have published presently accepted species names, only one third the number of authors (20) have published infrasubspecific names, even though there are nearly four times as many infrasubspecific names. Indeed, just three of these authors (Bruno Pittioni, Edgar Kriiger and Alexander Skorikov) are responsible for 70% of the infra- subspecific names (all of the species with many infrasubspecific names had been described before these three authors became active in publishing infra- subspecific names between 1910 and 1960, see Fig. 7). Many similar examples are known from work on butterflies (R. I. Vane-Wright, pers. com.), with au- thors choosing a particular favoured species and describing large numbers of infrasubspecific names (e.g. Bright & Leeds, 1938). The third point is that the three most prolific authors all worked in Europe, and there is a correlation among all 239 bumble bee spe- cies between the number of infrasubpecific names and the breadth of the species’ distributions just within Europe (measured as the number of occupied 611,000 km? grid cells between Britain and the Urals, but excluding Atlantic islands, North Africa, Turkey and the Caucasus; Spearman r= 0.67,1,,,= 13.99,p< 0.001). Thus, a high proportion of the many infrasubspecific names were published by very few European authors, for previously described species that are also particu- larly widespread in Europe. High numbers of synonyms and infrasubspecific names for B. terrestris and B. lucorum (subgenus Bombus) and for B. humilis and B. pascuorum (subgenus Thoracobombus) in Fig. 6 raise the possi- bility that large numbers of names are associated with particular groups of species, perhaps with particular subgenera. Number of names per species is plotted against range size per species for subgenera in Fig. 8. These properties are correlated (log-transformed data, correlation r= 0.58, F, ,.= 18.16, p< 0.001), but it is the subgenera with high scores that are more informative. The subgenus Kallobombus includes many names, but only a single, very variable species B. soroeensis, which is broadly distributed in Europe (see below and Reinig, 1939: fig. 10). The subgenera Alpinobombus and Laesobombus also have broadly distributed 86 P.H. WILLIAMS Table 2 Results of multiple regression of numbers of infrasubspecific names (from a manuscript catalogue, unpublished) on date of first formal description for presently accepted species and range size (number of occupied 611,000 km? grid cells world-wide). Species without infrasubspecific names were excluded from the analysis. Partial r values indicate the correlations with the infrasubspecific names variable after adjusting for the other predictor variable in each case. log Ginfrasubspecifics+1) = 14.169(+3.638) — 0.007(+0.002).date + 0.742(+0.212).log, (range) multiple r= 0.66 partial r date —0.364 log, (range) 0.340 eee 36:57 p< 0.0001 to4 P -3.79 < 0.001 3.50 < 0.001 20 Ne — 140 120 100 7) ® 5 By 80 L Oo ® o Te) 60 >| @ = rupestris = ruderarius 40 s pomorum lapponicus pratorum 20 0) 10 20 30 lucorum s.| a terrestris A soroeensis a humilis A pascuorum a 40 50 60 70 80 Synonyms Fig.6 Scatterplot of 239 presently accepted bumble bee species by numbers of infrasubspecific names and numbers of synonyms/subspecific names (from a manuscript catalogue, unpublished). The British fauna is distinguished as filled triangles, the nearly circumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some widespread European species are labelled individually. species but relatively few names, perhaps because they are absent or not abundant in those parts of Europe where the authors publishing most bumble bee names have worked, despite several of the species being very variable in colour pattern (e.g. B. balteatus). In con- trast, the high ratio of names per species for the subgenus Bombus shows the keen interest by some European authors such as Kriiger (1951, 1954, 1956, 1958) in describing the finer points of variation, not so much within the North American species, but particu- larly within the widespread European species, B. terrestris and B. lucorum. Summary of historical and regional trends in describing bumble bees Based on the evidence of asymptotic tendencies in species-discovery curves, a higher proportion of all CHECKLIST OF BUMBLE BEES 140 120 100 n oO § S 80 fc) i (Ss) oO a a 60 =I Y . © soroeensis = rupesins = a humilis Ta ruderanus fa 40 a pascuorum pomorum lapponicus © a consobrinus a Se as 1750 1775 1800 1825 87 1875 keriensis asiaticus O hypocrita 1900 1925 1950 1975 2000 Date of first description of species Fig. 7 Scatterplot of 239 presently accepted bumble bee species by numbers of infrasubspecific names (from a manuscript catalogue, unpublished) and date of first formal description of species. The British fauna is distinguished as filled triangles, the nearly cireumpolar fauna (B. hyperboreus, B. balteatus, B. polaris and B. lapponicus) as squares, and some species with more infrasubspecific names are labelled individually. species appear to be known for bumble bees than for many other groups of organisms. Most of these bum- ble bee species have been described by authors working in Europe (including European Russia). The species with the largest geographic range sizes, and particu- larly the European species with the largest ranges, have tended to be described first. The same species have also attracted the highest numbers of synonyms and subspecific names. As a group, bumble bees have an unusually high ratio of synonyms and subspecific names per species, which is otherwise known for some of the groups of larger and more colourful butterflies. A few European authors were disproportionately prolific between 1910 and 1960 in describing finer variation at infrasubspecific rank, which now accounts for one third of all bumble bee names. Again, this more detailed effort has been largely concentrated on the earlier-described species that are more widespread within Europe (in contrast, New World bumble bees have been ignored at this level), presumably because large samples were more readily accessible to the most active authors. Determining whether this re-direction of activity towards lower nomenclatural ranks was a | logical progression in the recognition of useful taxa, a | fashion in taxonomic concepts, or in some cases merely a less disruptive channelling of the enthusiasm of some authors to publish more names (the ‘mihi itch’), is beyond the scope of this preliminary review. All of these patterns in the descriptions of bumble bees must, as yet, be interpreted with caution. Much work still remains to be done on the rates of descrip- tion of taxa at different nomenclatural ranks (species, subspecies, infrasubspecies), on rates of recognition of synonymy and of changes in rank, and particularly on how this activity is partitioned among different time periods, different geographic regions, different taxo- nomic subgroups and different authors. Fundamental to almost all analyses are taxonomic revisions and checklists of bumble bee species. A revised checklist is now overdue, because nearly half (49%) of all names for bumble bees have been pub- lished since the last synoptic checklist (Skorikov, 1922a). Development of a revised checklist To begin to bring a checklist up to date, a draft was made in 1980 and first circulated for comment in 1985 (Williams, 1985a). This project was developed during a more detailed study of the west Himalayan fauna (Williams, 1991) and as part of continuing work on the large fauna of China in collaboration with Wang S.-f. 88 Kallobombus a Median names per species Confusibombus fo) Thoracobombus Bombus s.str a 0 10 20 30 P.H. WILLIAMS Laesobombus O° Alpinobombus ° 40 50 60 70 Median range size per species Fig. 8 Scatterplot of 38 bumble bee subgenera by median numbers of all names per species (including synonyms, subspecific and infrasubspecific names; from a manuscript catalogue, unpublished) and median range size per species (number of occupied 611,000 km? grid cells world-wide). The subgenera represented in the British fauna are distinguished as filled triangles and some subgenera are labelled individually. and Yao J. (unpub.). Some of the broader revisions that have had the greatest influence on this include works by Vogt (1909, 1911), Franklin (1913), Stephen (1957), Milliron (19706, 1971, 1973a, b), Loken (1973, 1984), Pekkarinen (1979), Reinig (1981), Wang (1982, 1987, 1988), Rasmont (1983, 1988), Thorp ef al. (1983), Labougle (1990), and especially the publications by Skorikov (1910-1938) and Tkalcti (1959-1989). In- evitably, the present checklist cannot be expected to solve all biological and nomenclatural problems, but it is hoped that by identifying some of the major prob- lems it will stimulate further research. Acknowledgements My grateful thanks to all who have contributed to the discussion of this and previous lists, including Donald Baker, Andreas Bertsch, Barry Bolton, Sydney Cameron, Gabriela Chavarria, Sally Corbet, Liz Day, Mick Day, Anne Divers, George Else, Kevin Gaston, Chris Humphries, Ian Kitching, Astrid Loken, Rod Macfarlane, Jim Mallet, Russell Miller, Chris O’ Toole, Antti Pekkarinen, Chris Plowright, Oliver Prys-Jones, Robert Prys-Jones, Pierre Rasmont, Malcolm Scoble, Chris Starr, Bill Stephen, Robin Thorp, Borek Tkalcii, Dick Vane-Wright, Doug Yanega, Yao Jian and Wang Shu-fang, although they do not necessarily share the opinions expressed here. I particularly appreciate the help of Philip Tubbs, Executive Secretary to the ICZN, for advice on the application of the current Code (ICZN, 1985) to nomenclatural problems. My thanks to Julie Harvey and Lorna Mitchell of the Entomology Library (Dept of Library and Information Services, NHM) for all their help. I would also like to thank Wang Shu-fang, Chen Wei and Yao Jian for their generous hospitality during my visit to China, as well as the Dept of Botany (NHM) for funding the visit. TAXONOMY Bumble bees are a monophyletic group (Williams, 1985b, 1995), constituting the tribe Bombini. They may be distinguished from other bees (family Apidae) by the following diagnosis (from Williams, 1991, which includes descriptions of the characters and dis- cussion of homologies): Bombini have the labrum at least twice as broad as long. The CHECKLIST OF BUMBLE BEES labrum lacks a longitudinal median ridge, although for the females it has a strong transverse basal depression. The clypeus has a transverse subapical depression and the apico- lateral corners are curved back towards the occiput. A malar area (= malar space) separates the compound eye from the base of the mandible, often by a distance greater than the breadth of the mandible at its base. The hind wings lack a jugal lobe (= anal lobe). The volsella (= lacinia) of the male genitalia is greatly enlarged and is produced apically beyond the gonostylus (= squama). Bumble bees are large (body length 7-27 mm) robust insects. Their bodies have a dense covering of variously coloured long plumose hairs, although these are few or absent on some parts of the ventral surface of the gaster, on parts of the propodeum, on parts of of the anterior face of gastral tergum I, and on parts of the head. The sclerites are usually black, or lighter brown on the distal parts of the limbs, but are never marked with bright yellow, red or metallic (= interfer- ence) colours. The wings may be transparent (= hyaline) to strongly darkened (= infuscated), but rarely show strongly metallic reflections. Female bumble bees have 12 antennal ‘segments’ (= scape, pedicel and 10 flagellomeres) and six visible gastral terga and sterna (abbreviated to TI-VI, SI-VI). Males have 13 antennal ‘segments’ (= scape, pedicel and 11 flagellomeres) and seven visible gastral terga and sterna (abbreviated to TI-VII, SI- VID). Where possible, a divisive, ‘top-down’ approach to the description of bumble bee diversity has been fol- lowed, in the sense of concentrating initially on higher-rank relationships and then distinguishing pro- gressively the species groups, species and then variation within species (as opposed to beginning with de- scribed infraspecific taxa and searching ‘upwards’ for close relatives). At the rank of species, this accepts those putative species or species complexes that are supported by consistent evidence for separate status, and which can be reliably identified throughout their range for the purpose of mapping distributions. This kind of broad over-view at least has the potential to apply consistent criteria across all taxa, even though it is appreciated that not all taxa at the rank of species are necessarily of the same kind (Ackery & Vane-Wright, 1984; de Queiroz & Donoghue, 1988). Specialists will need to modify this list as further information becomes available for particular species groups. Phylogeny, supra-specific taxa and ordering of species From available cladistic evidence (Williams, 1991, 1995), use of Psithyrus as a genus for the social parasites separate from the remainder of the social bumble bees in Bombus can no longer be justified, so a single genus Bombus is used for all of the species of bumble bees (see the comments under the subgenus Psithyrus). A system of subgenera has become widely used by 89 specialists who wish to label assemblages of the more closely similar species. This system is summarised with subgeneric diagnoses and keys by Richards (1968). For a review of supraspecific classifications of bumble bees, see Ito (1985). The subgeneric system would be more useful if the names were applied only to strictly monophyletic groups. Unfortunately, Richards’s (1968) concepts of the bumble bee subgenera do not always agree well with recent estimates of phylogeny, because some of these assemblages now appear to be paraphyletic (e.g. Mendacibombus) or even polyphyletic (e.g. Sibiricobombus in the sense of Richards, 1968, in- cludes Obertobombus, whereas he placed B. (Sibiricobombus) flaviventris in Subterraneobombus) (Williams, 1991). Furthermore, the system of subgenera would prob- ably be more useful if it were simplified (e.g. Menke & Carpenter, 1984; and reply by Williams, 1985c). For example, in the New World, both the monophyletic fraternus-group of subgenera and the subgenus Fervidobombus are endemic, and these are the only two groups represented south of the Panama isthmus. But whereas Fervidobombus has been treated nearly consistently as a single, relatively large subgenus (20 species in this list), the fraternus-group (18 species in this list) has regularly been split into as many as nine subgenera. However, no attempt is made in this checklist to revise radically the subgeneric system, because stabil- ity will only be served when a revision can be supported by a comprehensive cladistic analysis. This should include not only a broad sample of species, but also a broad range of morphological and molecular charac- ters. Minor modifications from the subgeneric system described by Richards (1968) are detailed in the list after the subgeneric names. Full synonymy of supraspecific names is included in this checklist, along with details of type species, because these have been revised since Richards (1968). The given generic combination for subgeneric names is Shown. Where a genus-group name was published at the rank of genus and subsequently treated at subgeneric rank, the first such action is listed separately. The two- letter abbreviations for subgeneric names are based on those used by Ito (1985). Species are listed in an order (Table 3) that repre- sents their phylogenetic relationships (after the sequencing convention of Nelson, 1972) as these are currently understood from cladistic studies of the adult morphology of both sexes (Williams, 1995, and many references therein). Within subgenera, this informa- tion is still of a very preliminary nature (e.g. Williams, 1991). Many other estimates of relationship exist and would result in different sequences of species names. An alphabetic index is provided as an aid to finding names in this list. 90 Table 3 List of names for subgenera of the genus Bombus, with numbers of species recognised in this checklist. The subgeneric classification is based on Richards (1968), modified to accommodate recent publications (see text; no attempt is made to revise the subgeneric system, because stability will only be served when a revision can be supported by a comprehensive cladistic analysis). Subgenera are listed in an order that represents their phylogenetic relationships (after the sequencing convention of Nelson, 1972) as these are currently understood from cladistic studies of the adult morphology of both sexes (Williams, 1995). Subgenus Number of species 1 Mendacibombus 12 2, Bombias 2 3 Confusibombus 1 4 Mucidobombus 1 5 Eversmannibombus 1 6 Psithyrus 29 7 Laesobombus 1 8 Orientalibombus 3 9 Exilobombus 1 10 Thoracobombus 19 11 Tricornibombus 3 12 Fervidobombus 20 13 Senexibombus 4 14 Diversobombus 4 15 Megabombus 14 16 Rhodobombus 3 17 Kallobombus 1 18 Alpinobombus 3) 19 Subterraneobombus 9 20 Alpigenobombus 6 2] Pyrobombus 43 22 Festivobombus 1 23 Rufipedibombus 2 24 Pressibombus 1 5 Bombus s.str. 10 26 Cullumanobombus 4 27 Obertobombus 2, 28 Melanobombus 14 29 Sibiricobombus 5 30 Fraternobombus 1 31 Crotchtibombus 1 32 Robustobombus 3) 33 Separatobombus 2 34 Funebribombus 2} 35 Brachycephalibombus 2 36 Rubicundobombus 1 37 Coccineobombus 2 38 Dasybombus 2 Criteria to discriminate species It is not possible or appropriate to discuss species concepts in detail in this paper (though the selected references provide some introduction; for recent re- views, see Claridge et al., 1997; Mallet, 1997). However, in order to interpret the checklist, where possible it would be useful to make the species-dis- criminating criteria explicit. It is equally important to convey the present belief that there is no simple solu- P.H. WILLIAMS tion to the problem, and that no single known approach can resolve all of the cases in a uniform and entirely satisfactory manner. Species concepts (ideas or general notions of the class of objects) and species diagnoses (operational determinations of individual objects) are contentious and probably unresolvable issues. Therefore there is arguably no single ‘true’ list of species, only more or less valid interpretations from different viewpoints. Unresolveable conflicts may arise from opposing views of the nature of species. Species have been regarded either as typological classes, with member- ship to be defined by some shared essence (reviewed by Templeton, 1981), or as individuals, to be discoy- ered (Ghiselin, 1975). There are also conflicting opinions concerning criteria (characteristics or stand- ards by which an object may be judged) for recognising species, based in part on differing emphasis on pattern or process (de Queiroz & Donoghue, 1988). Species may be considered not to differ from taxa at other ranks (e.g. genera, subspecies) in any qualitative way. There may be quantitative differences in the numbers of character differences that distinguish them in comparison with taxa of lower rank. For example, according to Mallet (1995:294), Darwin (1859) held this view. The problem with quantitative criteria (whether applied to genetic or phenotypic characters) is there is no reason to believe that any choice of threshold in the degree of difference used to recognise taxa at the rank of species is anything other than essentially arbitrary and thus idiosyncratic to particu- lar authors. In another view, species may be considered to differ qualitatively from taxa at other ranks. It is widely accepted, though often implicitly, that taxa at the rank of species should be recognised so as to mark the boundary between, on the one hand, reticulate rela- tions (for sexually reproducing organisms), and on the other, more consistently divergent genealogical rela- tions. One problem is that this distinction may require predictions as to whether or not currently distinct groups of individuals are likely to show reticulate relationships again in the future. Interbreeding and the associated genetic recombi- nation is an important part of Dobzhansky’s (1937) ‘modern synthesis’ of Mendelian genetics with Dar- win’s natural selection theory for evolution. Emphasising interbreeding as a criterion for recognis- ing species characterised what Mayr (1940, 1963) called the ‘biological’ species concept. These ideas have been modified in the recognition concept of species (Paterson, 1985). One problem with inter- breeding or mate recognition as criteria for recognising species is that direct and reliable evidence is rarely available and the results of tests under artificial condi- tions cannot necessarily be generalised (Splitter, 1982). Another is that the capacity for interbreeding is an ' | ——eE CHECKLIST OF BUMBLE BEES ancestral condition (i.e. not an homology) and so cannot provide support for recognising taxa in the phylogenetic sense (Rosen, 1979). In practice, all that is usually available to discrimi- nate species as ‘different’ is evidence from character differences and their patterns of concordance among individuals. The phylogenetic species concept (Cracraft, 1989) is popular because it also embodies the notion that species mark the boundary between different patterns of relationship among individuals and yet it does not rely on inference of interbreeding. The problem is that discovery of phylogenetic species as minimum cladistically-diagnosable (discrete) groups of individuals requires that these groups uniquely share homologies (synapomorphies), which may not always be the case (Ackery & Vane-Wright, 1984; Frost & Kluge, 1994). Mallet (1995) has argued for minimising the number of assumptions built into species concepts. He sug- gests that two nominal taxa should be considered conspecific until it can be demonstrated that data for multiple characters distinguish consistent subgroups of individuals with few or no intermediates (the char- acter-cluster concept of species). Although he was arguing against the use of the widely-held biological species concept, he recognised that his prescription differs little from recent common practice. The prob- lem with the cluster concept is how to decide on a threshold for permissable numbers of intermediate individuals between taxa for them still to be consid- ered separate species. Ultimately, species may be seen as useful conven- tions to aid in the communication of information gathered about the individuals that are their parts. It may be argued that the most important initial goal is to describe the nature of the variation in each particular case and to avoid presenting only theory-laden (and constrained) interpretations. In this way, basic infor- mation on variation will remain available for re-interpretation as theory changes. For the sake of illustration, four principal classes of problems in geographical variation may be distin- guished within the spectrum of kinds of relationships, with the following examples: Broad co-occurrence of differing individuals Skorikov (1931) and Reinig (1935) recognised that throughout much of the range of B. keriensis, both yellow-banded and cream- or white-banded individu- als with indistinguishable morphology co-occur (Fig. 9). From available evidence, it is possible that B. niveatus / vorticosus may show a similar pattern of _ yellow/white variation, as may B. impetuosus/potanini, although with differing degrees of geographical varia- tion in colour-form frequency (see the comments on these species). Consequently, taxa in these pairs are 91 also treated as conspecific for the present (it is possible that in some cases such colour differences may be controlled by alleles at a single locus, see Owen & Plowright, 1980, on B. melanopygus; and Williams, 1991, on B. asiaticus; or by small numbers of loci, see Plowright & Owen, 1980, on B. rufocinctus). In con- trast, although the yellow-banded B. shaposhnikovi and the white-banded B. handlirschianus also show a broadly-overlapping pattern of distribution, the one white-banded male that I have seen is distinct from the yellow-banded males in the morphology of its genita- lia (Williams, 1991). Broad clinal variation Many species show broad trends in variation across continents, most obviously in colour pattern (e.g. B. cingulatus, Fig. 10; and the trifasciatus-group, Fig. 13, which may be combined with locally convergent colour variation, e.g. within the haemorrhoidalis- group, breviceps-group and rotundiceps-group, see Sakagami & Yoshikawa, 1961; Tkalcii, 19685, 1989). In North America, several pairs of nominal taxa were described originally from individual type-specimens with differing colour patterns from eastern and west- ern regions respectively (e.g. B. auricomus /nevadensis, B. fervidus / californicus, B. pensylvanicus / sonorus, B. terricola / occidentalis). These taxon pairs have long caused difficulties, for example with Franklin (1913:239) commenting on a list including these taxa and others that are now considered conspecific that “it must be entirely a matter of personal opinion whether they should be given full species rank or be considered as only subspecies’ (although, intriguingly, B. auricomus /nevadensis were not included in Franklin’s list). In at least some of these cases, many individuals with what appears to be a continuum of intermediate colour patterns are now known from broad intervening areas, so that threshold criteria for distinguishing these taxa appear to be essentially arbitrary (e.g. making decisions based on whether a particular tergum has the pubescence entirely yellow, rather than having a few black hairs present). In consequence, taxa in these taxon pairs are treated here as conspecific and maps are compiled for the more clearly recognisable, more inclusive taxa (but see the comments on B. auricomus / nevadensis). Narrow hybrid zones In some cases, otherwise discrete colour forms with closely similar morphology meet in narrow zones (of the order of a few km in breadth), where there may be evidence of intermediate or genetically recombinant individuals. In Europe this is best known for B. ruderatus / argillaceus (Fig. 11; Scholl, Obrecht & Zimmermann, 1992), and inAsia it has been suggested oD P.H. WILLIAMS | § ‘ Fig.9 Approximate distribution range (area within the dotted line) and principal colour variation for B. keriensis from Reinig (1939: fig. 23). Many more records are available now, but the pattern remains similar, with broad overlap of yellow- and white-banded individuals in Mongolia, Tien Shan, Pamir and western Himalaya. Yellow and cream pubescence is shown on the bees by crosses; red pubescence by vertical hatching. Fig. 10 Distribution records (spots), approximate range (area within the line) and principal colour variation for B. cingulatus in the northern Palaearctic Region from Reinig (1939: fig. 7). The lightest individuals occur in the east (Kamchatka) and the darkest individuals (with the black thoracic band) occur in the west, with intermediate individuals in intervening areas. CHECKLIST OF BUMBLE BEES 93 ombus : /uderans suderatis abr OTGUMACEUS SCOP. Fig. 11 Distribution records (spots), approximate range (area within the cross-hatching, left, and line, right) and principal colour variation between queens of B. ruderatus and B. argillaceus in Europe from Reinig (1939: fig. 7). These taxa were regarded as subspecies by Reinig, but have recently been treated as separate species. Although there is evidence of a hybrid zone between some areas of parapatry, the hybrid individuals are very rare (Scholl, Obrecht & Zimmermann, 1992). Yellow pubescence is shown on the bees by crosses. my Fig. 12 Distribution records (spots) and principal colour variation for B. asiaticus in Kashmir from Williams (1991: map 48). There is evidence of a hybrid zone between some areas of parapatry, such as some high passes along the divide of the Great Himalaya Range, where there are abundant hybrid individuals. The spot symbols show the locally most abundant colour pattern. Yellow pubescence is shown on the bees by fine stippling; red pubescence by vertical hatching. 94 P.H. WILLIAMS Fig. 13 Distribution records and principal colour variation within the trifasciatus-group in Asia (updated from Williams, 1991: fig. 11). The individuals may all be considered parts of a single species, B. trifasciatus, depending on which species-defining criterion is accepted. The dashed line shows the 1000 m contour above sea level and the solid line shows the 4000 m contour. Yellow pubescence is shown on the bees by fine stippling, orange pubescence by coarse stippling, red pubescence by vertical hatching. CHECKLIST OF BUMBLE BEES « On | in On <@ 96 for B. asiaticus / longiceps (Fig. 12; Williams, 1991). Other possible examples include B. lapponicus / monticola in Europe and B. pyrosoma / friseanus / miniatus in China. In the case of B. asiaticus / longiceps, | have treated them as conspecific, because intermediate individuals greatly outnumber ‘typical’ individuals at some localities. For the other cases, I have followed earlier treat- ments of these taxa as separate species, because intermediate individuals are rare or not well known (although this may be a consequence of poor sam- pling in some inaccessible areas). Disjunct peripheral populations Some peripheral populations on offshore islands or habitat islands (e.g. mountains) show some diver- gence in colour pattern with little morphological divergence. European examples include B. terrestris / canariensis and B. hortorum / reinigiellus. Asian examples include B. schrencki / honshuensis, B. trifasciatus / maxwelli (Fig. 13, Peninsular Malay- sia), B. trifasciatus / wilemani (Fig. 13, Taiwan), B. breviceps / angustus, B. parthenius / sonani, B. flavescens / rufoflavus and B. flavescens / baguionensis. For the application of the biological species concept, in these cases there is no ‘natural’ meeting of individuals between the taxon pairs and so no admissible evidence on interbreeding (Splitter, 1982). For the application of Mallet’s (1995) cluster concept, quantitative analysis of patterns of variation is urgently needed. Where this information is absent, I agree with his prescription of treating taxa in these taxon pairs as provisionally conspecific. Bombus honshuensis and B. schrencki are mapped separately here because, from published accounts and a small sample of material examined, their colour differ- ences appear to coincide with stronger and more consistent morphological distinctions. It is hoped that further information may help to clarify these cases. In the interests of pluralism, I aim to report not only a preferred interpretation in the comments on each species, but also at least the more widely-held alternative interpretations. Sub-specific taxa For this checklist the interest is primarily in problems of recognition and nomenclature for taxa at the rank of species. Subspecific names refer to parts of species, and so for present purposes these can be treated as synonyms of specific names (e.g. Schwarz ef al., 1996). This is not to say that subspecific taxa should not be recognised if they are considered useful, and of course other biologists may add subspecies to this list (cf. Rasmont et al., 1995). P.H. WILLIAMS NOMENCLATURE Nomenclature should be seen as the servant of biol- ogy: its purpose is to provide labels that enable biologists to communicate information about organ- isms with minimal confusion concerning the organisms to which they refer. Accounts of the history of nomen- clature for many groups of organisms (e.g. on British bumble bees: Alford, 1975; Prys-Jones & Corbet, 1987:82) show that this is not a trivial matter and that rules are necessary. Treatment of names follows the /nternational Code of Zoological Nomenclature (International Commis- sion on Zoological Nomenclature [{CZN], 1985). The Principle of Priority is generally adhered to, although regard is given to the stated purpose of priority (ICZN, 1985: Article 23b): namely that it should be used to promote stability and is not intended to be used to upset a long-accepted name in its accustomed mean- ing (Article 79c) through the introduction of an unused name that is its senior synonym. Similar action is also suggested where cases of homonymy affect current usage, although this action cannot be taken when it is felt desirable to maintain availability of a senior homo- nym. My suggestions for applications to ICZN for conservation of names in current use are indicated by stars (@). Typographical conventions Bombus b-us valid name in the species group, c-us available name in the species group, including synonyms of a valid spe- cies name, 2d-us available name in the species group, a provisional synonym of a valid species name, unavailable name, informally asso- ciated with a valid species name, jf-us examined type material for species-group name f-us examined (in whole or in part), [e-us] @ comments on status of species, O comments on application of names, Ok) suggestion for application to ICZN. ?Bombus g-us_ valid name in the species group, fora taxon that is recognised provision- ally as a separate species from B. b-us. A question mark (?) before a valid name shows that, while it refers to a taxon that is considered likely to be a separate species, it may be conspecific with the preceding taxon in the list (1.e. while Bombus g-us may be conspecific with Bombus b-us, Bombus d-us is much more likely to be conspecific with Bombus b- US). CHECKLIST OF BUMBLE BEES Names in the more detailed references are followed by names of authors, date of first publication (within the meaning of ICZN, 1985), and page reference. Wherever possible, the true first date of publication is given in preference to any purported date of publica- tion when these differ. If a name were published originally in a different generic combination, then the original genus is shown in brackets. If the name had been published originally with a different termination, or with capital initial letters, diacritic marks etc., then the original form is shown without the mandatory changes (with the exception that small capital letters are reduced to lower case). Selection of synonyms This checklist is based on a much longer catalogue of over 2800 names. As a checklist, it is not required to include the full list of synonyms, so synonyms are selected for this list primarily where they help to clarify the identity and scope of the species (including the subspecies included by some authors), particularly with reference to those names in most common use in the literature of the last 25 years. Misidentifications are not included with the lists of synonyms and are discussed only when necessary to clarify the applica- tion of problematic names. Applications to ICZN Flexibility in interpretation of the status of taxa is possible where the evidence to distinguish among interpretations is absent, inconclusive, or may permit different interpretations under different species con- cepts. Otherwise flexibility in the application of names depends on whether systematists are eager to apply to the International Commission on Zoological Nomen- clature to use its Plenary Power in order to conserve a preferred usage of names (e.g. Laken et al., 1994; ICZN, 1996). I propose that this could be achieved in a single application to include all names for which action is currently known to be required (atratus, balteatus, distinguendus, flavifrons, humilis, hyperboreus, mesomelas, mixtus, norvegicus, polaris, pyrenaeus, soroeensis and variabilis). Comments on this proposal would be welcomed. DISTRIBUTION MAPS This checklist was compiled in conjunction with dis- tribution data in support of biogeographic studies. 97 Maps of world-wide distribution at a coarse grain size were designed for use in comparisons of regional bumble bee faunas (e.g. Williams, 1989, 1991, 1993, 1995, 1996a, b; Williams & Seddon, 1993; Williams & Humphries, 1996). Aside from any difficulties in identifying species or localities, comparisons among faunas are complicated by two principal factors: first, by differences in sam- pling effort (as illustrated by ‘species-accumulation curves’, e.g. Colwell & Coddington, 1994); and sec- ond, by differences in the extent of sampling areas (‘species-area effects’, e.g. Connor & McCoy, 1979). Fortunately for the first problem, the attractiveness of bumble bees to collectors has ensured that they have been relatively intensively sampled, so that most fau- nas are relatively well known. But in order to reduce this problem further, rather than extrapolate local rich- ness and lose information on individual species, the expected distributions of some species are interpo- lated on the basis of knowledge of their habitat associations (see the legend to Fig. 14). To reduce the second problem of species-area effects, equal-area grid cells were established using a cylindrical, equal- area projection of the world, marked at intervals of 10° of longitude and calculated intervals of latitude (Fig. 14). However, this does not ensure equal land areas among grid cells, or equal areas of habitat suitable for bumble bees. Because the intention is to study biogeographic patterns, maps are required to show all historical records, including data from areas where species may now be extinct. On the other hand, data exclude fossil taxa (reviewed by Zeuner & Manning, 1976) and documented introductions (e.g. Oliff, 1895; Frison, 1925b; Gurr, 1957; Prys-Jones et al., 1981; Arretz & Macfarlane, 1982; Cardale, 1993). The maps for every species are not included with this checklist because many data are still being col- lected, although for each subgenus a preliminary map of species richness is included as a general guide (or for monotypic subgenera, a map of records for the single species is included). The numerical values for the grey-scale classes differ between maps and are not shown. This is because I have adopted an alterna- tive approach of using equal frequency classes, which have the advantage that each grey-scale class remains consistent in its relative richness among all maps (e.g. dark grey always shows the richest one fifth of occu- pied cells excluding the maximum etc.). The sources of the distribution data have not been included be- cause this will be included in a later atlas. 98 P.H. WILLIAMS Pp Key to map symbols: Maps for single species specimens examined, precise literature records (e.g. ‘Dungeness TRO1, UK’), vague locality data (e.g. “Florida’), interpolations of expected distribution (following common practice for range-filling maps; the rules adopted here are to fill cells between occupied cells when filled cells are known to have had a high proportion of suitable habitat within recorded history; these records amount to < 10% of all gridcell records at this scale, Williams, 1993). OOS®e Maps for multiple species maximum species counts are shown in black, otherwise counts are divided into five grey-scale classes of approximately equal size by numbers of grid cells. Fig. 14 Map of the world (excluding Antarctica) using a cylindrical equal-area projection that is orthomorphic (minimum shape distortion) at 46° North and South (where bumble bee records are particularly plentiful). Intervals of 10° longitude (top of map) are used to calculate intervals of latitude (right of map) that provide equal-area grid cells of c. 611,000 km?. The portion of the grid shown covers the known, native distribution of bumble bees. Map symbols are shown above for (a) plotting individual species, for which different spots distinguish different data categories (Map 3); or (b) for plotting coincidence maps for multiple species, using a grey scale for variation in species richness (Map 1). CHECKLIST OF BUMBLE BEES LIST OF SPECIES (plot of total species richness with grey scale, for explanation see Fig. 14) Genus BOMBUS Latreille in the broad sense [Bremus [Jurine], 1801:164, type-species Apis terrestris Linnaeus (= Bombus terrestris (Linnaeus)) by subse- quent designation of Morice & Durrant, 1915:429, suppressed by ICZN, 1939] Bombus Latreille, 1802a:437, type-species Apis terrestris Linnaeus (cited asApis terrestris F.) (=Bombus terrestris (Linnaeus)) by monotypy Bombus Latreille, 1802b:385, type-species Apis terrestris Linnaeus (=Bombus terrestris (Linnaeus)) by monotypy, redescribed [Bremus Panzer, 1805:pl. 19-21, type-speciesApis agrorum Fabricius (= Bombus pascuorum (Scopoli)) by subse- quent designation of Sandhouse, 1943:532, suppressed by ICZN, 1954] [Bombellus WE, 1931:248, incorrect subsequent spelling] Subgenus MENDACIBOMBUS Skorikov Mendacibombus Skorikov, 1914a:125, type-species Bombus mendax Gerstaecker by subsequent designa- tion of Sandhouse, 1943:572 Bombus (Mendacibombus) Kriiger, 1917:62 COMMENT. The species of Mendacibombus appear to be paraphyletic with respect to the rest of the oY) bumble bees and in consequence are not a ‘natural’ group (Williams, 1991, 1995). Bombus (Md.) avinoviellus (Skorikov) avinoviellus (Skorikov, 1914a:126 [Mendacibombus]) ex- amined callophenax Cockerell, 1917:122, examined Bombus (Md.) mendax Gerstaecker mendax Gerstaecker, 1869:323, examined latofasciatus Vogt, 1909:50, not of Vogt, 1909:42 (= B. lucorum (Linnaeus)) pyrenes (Tkalcit, 1975:173 [Mendacibombus]) replace- ment name for latofasciatus Vogt, 1909:50 Bombus (Md.) makarjini Skorikov makarjini Skorikoy, 1910a:329, examined Bombus (Md.) superbus (Tkalcit) superbus (Tkalctt, 1968a:22 |Mendacibombus]) examined Bombus (Md.) himalayanus (Skorikov) ?varius (Skorikoy, 1914a:125 [Mendacibombus]) exam- ined, not of Lepeletier, 1832:381 (= B. campestris (Panzer)) himalayanus (Skorikov, 1914a:127 [Mendacibombus}) examined Bombus (Md.) marussinus Skorikov marussinus Skorikov, 1910a:330, examined afghanus Reinig, 1940:230, examined Bombus (Md.) turkestanicus Skorikov turkestanicus Skorikoy, 1910a:329, examined Bombus (Md.) defector Skorikov defector Skorikov, 1910a:330 ?altaicus Skorikov, 1910a:329, not of Eversmann, 1846:436 (= B. melanurus Lepeletier) ?margreiteri Vogt, in Skorikov, 1910a:330, examined @ TAXONOMIC sTATUS. Skorikov’s (1910a) de- scriptions of varieties of B. mendax are all of females. Many of these nominal taxa have subsequently been treated as separate species (e.g. Skorikov, 1931; Rasmont, 1988). However, I have examined type material or other material identified by Skorikov for all of these taxa and find some of them to be morphologically closely similar. The females of defector, altaicus and margreiteri differ from one another principally in 100 colour, and the only males I have seen associated with them (collections in London, Petersburg, Beijing) have very similar genitalia (which are distinct from B. mendax). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall continue to treat B. defector, B. altaicus and B. margreiteri as parts of a single variable species, B. defector (Williams, 1985a, 1991). O NOMENCLATURE. Williams (1991) regarded B. defector, B. altaicus and B. margreiteri as likely to be conspecific and following the Principle of First Re- viser (ICZN, 1985: Article 24) chose B. defector as the name for the species. Bombus (Md.) handlirschianus Vogt Handlirschianus Vogt, 1909:49 Bombus (Md.) shaposhnikovi Skorikov shaposhnikovi Skorikoy, 1910a:329 Bombus (Md.) waltoni Cockerell chinensis Skorikov, 1910a:330, examined, not of Morawitz, 1890:352 (= B. chinensis (Morawitz)) waltoni Cockerell, 1910b:239, examined Bombus (Md.) convexus Wang lugubris Morawitz, 1880:339, examined, not of Kriechbaumer, 1870:159 (= B. maxillosus Klug) convexus Wang, 1979:190, examined Subgenus BOMBIAS Robertson Bombias Robertson, 1903:176, type-species Bombias auricomus Robertson (?= Bombus nevadensis Cresson) by original designation Bombus (Bombias) Franklin, 1913:138 Nevadensibombus Skorikoy, 1922a:149, type-species Bombus nevadensis Cresson by subsequent designation of Frison, 1927:64 Bremus (Boopobombus) Frison, 1927:59 (proposed as a P.H. WILLIAMS section name but stated by Frison to include those forms considered by Franklin, 1913, to belong to the subgenus Bombias Robertson), type-species Bombias auricomus Robertson (= Bombus auricomus (Robertson)) by sub- sequent designation of Williams, 1995:339. Bombus (Bi.) nevadensis Cresson nevadensis Cresson, 1874:102 COMMENT. A single queen of B. nevadensis has been reported from Hidalgo, Mexico, by Milliron (1971) and Hurd (1979), although the species is not listed for Mexico by Labougle (1990). ?Bombus (Bi.) auricomus (Robertson) auricomus (Robertson, 1903:176 [Bombias]) @ TAXONOMIC STATUS. B. nevadensis and B. auricomus have been regarded both as conspecific (e.g. LaBerge & Webb, 1962; Milliron, 1971; Thorp er al., 1983; Laverty & Harder, 1988) and as separate species (e.g. Franklin, 1913; Rasmont, 1988; Scholl, Thorp, Owen & Obrecht, 1992; Poole, 1996). B. nevadensis from western North America was not mentioned in the original description of B. auricomus (lectotype worker from Illinois by designation of Milliron, 1971:78), although the latter was described using characters of morphology and of colour pattern. The two taxa have generally been distinguished on the basis of the extent of the black pubescence on the dorsum of the female thorax and laterally on the male gastral terga (e.g. Franklin, 1913). The only study to investigate variation in characters used to distinguish the two taxa at a fine spatial scale in their area of overlap was by LaBerge & Webb (1962). They reported (p. 26) that “Throughout the broad middle half of Nebraska nevadensis seems to be rather rare and most specimens, although referable to sub- species auricomus show some indication of intergrading with the typical subspecies [nevadensis] in the west. . .. Many specimens from Nebraska in the range of the typical subspecies [nevadensis| show some tendency toward the darker coloration of subspe- cies auricomus. They concluded that these variable bees are all parts of the same species. Recently, Scholl et al. (1992) distinguished two groups of individuals on the basis of differing mobility morphs of five enzymes. The individuals in one en- zyme group were all extensively dark-banded, and Scholl et al. associated these with the name B. auricomus. However, individuals in the other enzyme group, which Scholl et al. associated with the name B. nevadensis, apparently included not only the contrast- ing, extensively pale individuals (B. nevadensis), but also a few of the extensively dark-banded individuals (B. auricomus) similar to those in the first group (8/49 CHECKLIST OF BUMBLE BEES individuals had gastral tergum I almost completely black; 3/49 individuals had the scutellum predomi- nantly black). Thus the enzyme evidence does identify two groups of individuals, but (1) these do not appear to correspond precisely to the two traditional colour groups; (2) some of the key areas likely to support intermediate or recombinant individuals still need to be sampled for enzyme variation (e.g. in the Dakotas, L. Day in litt.); and (3) inheritance of enzyme and colour states needs to be better understood, including the unusual enzyme morphs of the heterozygous bees (detected in 20/141 queens). They concluded that these bees represent two species. A. Scholl (in litt.) reports a further intriguing morphometric study. A random subsample of 20 queens from the enzyme study was scored for 15 characters and analysed by linear discriminant analysis. This method seeks a combination of characters that best discriminates any two a priori sets, in this case using three measurements of parts of the radial cell, eye and antenna. However, although this approach may be useful for discriminating previously recognised taxa, it does not provide evidence that they are necessarily separate species (it could also be used to discriminate morphological subsets within a single, variable popu- lation, e.g. among breeds of domestic dogs). From an examination of 41 females, so far I have found only one subtle morphological character to distinguish eastern, banded bees (B. auricomus), on the one hand, from western unbanded (B. nevadensis) and banded (e.g. Vancouver Island) bees, on the other. This concerns the anterior part of a band of large punctures along the inner eye margin, dorsally oppo- site the ocelli, just before these punctures meet a more anterior, very dense patch of small punctures. The western bees have areas between the large punctures conspicuously shining, with few fine punctures and lacking microsculpture. In contrast, the eastern bees have these areas appearing rather dull, often with more of the fine punctures, and more particularly with a very fine, wrinkled or reticulate microsculpture. A similar difference may be present in the males, posterio- laterally to the ocelli, though the sample sizes available to me are too small for much confidence. I regard the conflicting evidence available at present as not entirely conclusive as to whether these bees are parts of the same population or two separate species. As far as is known, both the variations of the colour pattern and of the enzyme mobilities are inherited and genetically determined, but details of patterns of in- heritance and of the spatial aspects of any association between these characters are unknown. In view of the multiple enzymes differences found and of the appar- ent association between the enzyme groups and the morphological character states, I shall follow the treat- ment of these taxa as two separate species until more evidence is available. (plot of records for a single species, for explanation and key see Fig. 14) Subgenus CONFUSIBOMBUS Ball Bombus (Confusibombus) Ball, 1914:78, type-species Bombus confusus Schenck by monotypy Bombus (Sulcobombus) Kriiger, 1917:65, type-species Bombus confusus Schenck by subsequent designation of Sandhouse, 1943:602 Confusobombus Skorikoy, 1922a:156, type-species Bombus confusus Schenck by subsequent designation of Richards, 1968:214 Bombus (Cf.) confusus Schenck confusus Schenck, 1859:135 paradoxus Dalla Torre, 1882:18 festivus Hoffer, 1882:80, not of Smith, 1861:152 (= B. festivus Smith) @ TAXONOMIC STATUS. B. confusus and B. paradoxus differ in the colour pattern of the pubes- cence (e.g. Reinig, 1939: fig. 19). Rasmont (1988) reports that in north western Europe, the yellow-banded and white-tailed B. paradoxus occurs only as rare individuals within the population of predominantly unbanded and red-tailed B. confusus. In contrast, all of the individuals that I have seen from the disjunct population in Central Asia have the yellow-banded and white-tailed B. paradoxus colour pattern. Subgenus MUCIDOBOMBUS Kriiger Mucidobombus Kriiger, 1920:350, type-species Bombus mucidus Gerstaecker by monotypy Bombus (Mucidobombus) Pittioni, 1937:97 102 Bombus (Mc.) mucidus Gerstaecker mucidus Gerstaecker, 1869:324 atratus Friese, 1911:572, examined Subgenus EVERSMANNIBOMBUS Skorikov Agribombus (Eversmannibombus) Skorikoy, 1938a:145, type-species Mucidobombus eversmanniellus (= Bombus persicus Radoszkowski) by monotypy Bombus (Eversmannibombus) Richards, 1968:214 Bombus (Ey.) persicus Radoszkowski calidus Eversmann, 1852:133, examined, not of Erichson in Middendorff, 1851:65 (= B. hypnorum (Linnaeus)) persicus Radoszkowski, 1881:v, examined Persicus Radoszkowski, 1883:214, redescribed eversmanni Friese, 1911:572, not of Skorikov, 1910¢:581 ( B. modestus Eversmann), replacement name for calidus Eversmann, 1852:133 eversmanniellus (Skorikoy, 1922a:149 [Mucidobombus]) replacement name for eversmanni Friese, 1911:572 Subgenus PSITHYRUS Lepeletier Psithyrus Lepeletier, 1832:373, type-species Apis rupestris Fabricius (= Bombus rupestris (Fabricius)) by subse- quent designation of Sandhouse, 1943:572 Apathus Newman, 1835:404, replacement name for Psithyrus Lepeletier, incorrectly stated to be a junior homonym of Psithyros Hiibner, [1819]:132 (= Macroglossum Scopoli, 1777:414) P.H. WILLIAMS ?Psithyrus (Laboriopsithyrus) Frison, 1927:69, type-spe- cies Bombus laboriosus Fabricius (= Emphoropsis laboriosus (Fabricius) in the sense of Frison (=Bombus citrinus (Smith), a misidentification, see Milliron, 1960:99, requiring designation by ICZN) by original fixation & Psithyrus (Ashtonipsithyrus) Frison, 1927:69, type-spe- cies Apathus ashtoni Cresson (= Bombus ashtoni (Cresson)) by original designation Psithyrus (Fernaldaepsithyrus ) Frison, 1927:70, type-spe- cies Psithyrus fernaldae Franklin (= Bombus fernaldae (Franklin)) by original designation Psithyrus (Eopsithyrus) Popov, 1931:134, type-species Apathus tibetanus Morawitz (= Bombus tibetanus (Morawitz)) by original designation Psithyrus (Metapsithyrus) Popov, 1931:135, type-species Apis campestris Panzer (= Bombus campestris (Pan- zer)) by original designation Psithyrus (Allopsithyrus) Popov, 1931:136, type-species Apis barbutella Kirby (= Bombus barbutellus (Kirby)) by original designation Psithyrus (Ceratopsithyrus) Pittioni, 1949:270, type-spe- cies Psithyrus klapperichi Pittioni (= Bombus cornutus (Frison)) by original designation Psithyrus (Citrinopsithyrus ) Thorp inThorp et al., 1983:50, type-speciesApathus citrinus Smith (=Bombus citrinus (Smith)) by original designation Bombus (Psithyrus) Williams, 1991:44 [Psithyrus (Fernaldepsithyrus) Amiet, 1996:86, incorrect subsequent spelling] @ TAXONOMIC sTATUS. It has long been consid- ered useful to regard Psithyrus as a separate genus in recognition of the distinctive behaviour of the species, as social parasites in colonies of the remaining Bombini, and in recognition of their distinctive mor- phology. However, most recent studies have shown (if phenograms are interpreted along with cladograms as phylogenetic estimates) that, although Psithyrus is itself very likely to be monophyletic, the remaining bumble bees are not (Plowright & Stephen, 1973; Obrecht & Scholl, 1981; Ito, 1985: Williams, 1985), 1991, 1995; Pamilo et al., 1987). I have previously attempted to retain the use of the names Psithyrus and Bombus for monophyletic genera by recognising a third genus, Mendacibombus (Williams, 1985b). However, further study of all of the species of Mendacibombus (Williams, 1991, 1995) showed that it is likely to be paraphyletic with respect to all other bumble bees, with the consequence that as many as another nine genera (mostly for single spe- cies) might be required to maintain monophyly alongside a genus Psithyrus. In the face of this evi- dence, a pragmatic solution was recommended, recognising a single genus Bombus for all bumble bees, to include Psithyrus as a subgenus. This is a return to an emphasis of the more widely shared characters and the more distant affinities for the ge- neric concept, encouraged by the opinion of Michener (1990) that bumble bees are ‘morphologically mo- CHECKLIST OF BUMBLE BEES notonous’ in comparison with variation among species within closely related groups such as Euglossini (or- chid bees) and Meliponini (stingless bees). One advantage of a single genus for all bumble bees is that it recognises a group for which evidence of monophyly is particularly strong, so that nomenclature is most likely to remain stable in the future. Use of a single genus Bombus for all bumble bees (Williams, 1991) has now been accepted by most recent authors (e.g. Rasmont & Adamski, 1995; Rasmont ef al., 1995; Schwarz et al., 1996). The subgenera within the former genus Psithyrus have often been considered less distinct from one another than have the other subgenera of Bombus (Pittioni, 1939a; Ito, 1985; Williams, 19855; Michener, 1990) and therefore may be treated as synonyms of Psithyrus (Milliron, 1961; Williams, 1991, 1995). In an alternative treatment, Rasmont et al. (1995) include the former subgenera of the former genus Psithyrus as separate subgenera within the genus Bombus. O NOMENCLATURE. The names of six species of the subgenus Psithyrus from Kashmir were explicitly stated to be new combinations with the genus Bombus by Williams (1991). Rasmont ef al. (1995) have since listed the other European species in this combination. No formal statements of new combination are made here for the remaining species of the subgenus Psithyrus because a principle of implied combinations (Poole, 1996) is followed after the change in status of Psithyrus from genus to a subgenus of Bombus. APPLICATION TO ICZN. Because the type spe- cies of Laboriopsithyrus was misidentified (discussed by Milliron, 1960:99), ICZN is required to designate as type species whichever species will best serve nomenclatural stability (ICZN, 1985: Art. 70b). It is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to select the species actually involved (Bombus laboriosus in the sense of Frison, = Bombus citrinus (Smith)), which was wrongly named in the type fixation (ICZN, 1985: Art. 70b(i)). COMMENT. The highest richness of species of the subgenus Psithyrus occurs in the Old World (there are no species known from south of Panama), al- though the earliest-diverging species appear to be North American (unpublished). This is the opposite pattern to that shown by species of the largest subgenus, Pyrobombus (see the comments on the subgenus Pyrobombus). All species of the subgenus Psithyrus are believed to be obligate social parasites in colonies of other Bombus species (reviewed by Alford, 1975; Fisher, 1987). There is variation in the degree of host specificity. See also the comments on B. inexspectatus and B. hyperboreus. 103 Bombus (Ps.) insularis (Smith) interruptus Greene, 1858:11, not of Lepeletier, 1832:381 (= B. rupestris (Fabricius)) insularis (Smith, 1861:155 [Apathus]) examined consultus (Franklin, 1913:459 [Psithyrus]) ? bicolor (Franklin, 1913:460 [Psithyrus]) not of H6ppner, 1897:33 (=B. soroeensis (Fabricius)) (provisional syno- nym) crawfordi (Franklin, 1913:464 [Psithyrus]) @ TAXONOMIC stTaTUS. According to D. Yanega (in litt.), who has examined the type material, B. bicolor Franklin is conspecific with B. interruptus. Bombus (Ps.) citrinus (Smith) citrinus (Smith, 1854:385 [Apathus]) examined contiguus (Cresson, 1863:112 [Apathus]) Bombus (Ps.) variabilis (Cresson) ® intrudens (Smith, 1861:154 [Apathus]) examined variabilis (Cresson, 1872:284 [Apathus]) new synonym ?guatemalensis (Cockerell, 1912:21 [Psithyrus}) (provi- sional synonym) ?sololensis (Franklin, 1915:173 [Psithyrus]) (provisional synonym) ?mysticus (Frison, 1925a:138 [Psithyrus]) (provisional synonym) @ TAXONOMIC STATUS. Specimens in the NHM collection from Mexico and Guatemala labelled ‘intrudens’ and ‘sololensis’ appear to me to be closely similar to B. variabilis. Frison (1925a) believed that B. sololensis is acolour form of B. guatemalensis. Never- theless, he proceeded to distinguish B. mysticus as a separate species on the basis of colour pattern alone. I am unaware of any reason (other than minor differ- ences in colour pattern) why B. variabilis, B. intrudens, B. sololensis, or B. guatemalensis and B. mysticus (judging from the published descriptions at least), should not be considered conspecific. O NOMENCLATURE. A female in the NHM collec- tion has three labels “Apathus / intrudens / Smith.’, *58.135 MEX. / (Oajaca.)’, “Holo- / type’ and I am unaware of any problems with this designation. If this is correct and the type is conspecific with B. variabilis, then B. intrudens is the oldest available name for this species. D. Yanega (in /itt.) agrees with this interpreta- tion. € APPLICATION TOICZN. Although B. intrudens is the oldest available name for the present interpretation of this species, the name B. variabilis has been in common use for the species since 1947 (e.g. Stevens, 1948; Chandler, 1950; LaBerge & Webb, 1962; Mitchell, 1962; Medler & Carney, 1963; Hobbs, 1966; Plowright & Stephen, 1973; Hurd, 1979; Husband et al., 1980; Michener, 1990; Poole, 1996). I know of no publications using the name B. intrudens since 1947. It 104 is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). However, the consequence of this action would be that B. intrudens would no longer be available for a species or for a subspecies of B. variabilis (Cresson). Bombus (Ps.) suckleyi Greene Suckleyi Greene, 1860:169 Bombus (Ps.) vestalis (Geoffroy) veftalis (Geoffroy in Fourcroy, 1785[see Hagen 1862:246]:450 [Apis]) Bombus (Ps.) perezi (Schulthess-Rechberg) perezi (Schulthess-Rechberg, 1886:275 [Psithyrus]) Bombus (Ps.) ashtoni (Cresson) Ashtoni (Cresson, 1864:42 [Apathus]) Bombus (Ps.) bohemicus Seidl nemorum (Fabricius, 1775:380 [Apis]) examined, not of Scopoli, 1763:307 (= B. subterraneus (Linnaeus)), not of Fabricius, 1775:382 (= B. distinguendus Morawitz) bohemicus Seidl, 1837:73 ?chinganicus (Reinig, 1936:8 [Psithyrus]) (provisional synonym) hedini (Bischoff, 1936:26 [Psithyrus]) not of Bischoff, 1936:15 (= B. hedini Bischoff) @ TAXONOMIC sTATUS. Iam unaware of any rea- son (other than the small body size of the holotype female and three paratype females of B. chinganicus) why B. bohemicus and B. chinganicus should not be considered conspecific. Consistent with this, body sizes do appear to vary considerably within British species of the subgenus Psithyrus, including B. bohemicus. Bombus (Ps.) coreanus (Yasumatsu) coreanus (Yasumatsu, 1934:399 [Psithyrus]) Bombus (Ps.) barbutellus (Kirby) Barbutella (Kirby, 1802:343 [Apis]) examined ?richardsi (Popov, 1931:150,190 [Psithyrus]) not of Frison, 1930:6 (= B. rufipes Lepeletier) 2icenti (Maa, 1948:34 [Psithyrus]) examined O NOMENCLATURE. Loken (1984) interpreted B. saltuum (Panzer, 1801) as conspecific with B. barbutellus. Consequently, B. saltuum would appear to be the oldest available name for this species. How- ever, Lgken made no further comment on this and used the name Psithyrus barbutellus (= B. barbutellus), possibly because she remained unsure of the identity of B. saltuum. In contrast, Warncke (1986) interpreted P.H. WILLIAMS B. saltuum as conspecific with B. subterraneus. See the comments on B. subterraneus. ?Bombus (Ps.) maxillosus Klug maxillosus Klug in Germar, 1817:269 lugubris (Kriechbaumer, 1870:159 [Psithyrus]) unicolor (Kriechbaumer, 1870:159 [Psithyrus]) mixta (Kriechbaumer, 1870:160 [Psithyrus]) Susterai (May, 1944:267 [Psithyrus]) not infrasubspecific after Tkalci, 1977:224 @ TAXONOMIC STATUS. As Rasmont (1988) notes, B. maxillosus 1s closely similar to B. barbutellus in morphology and habitat, so that specimens cannot always be distinguished reliably. Consequently these nominal taxa might be considered conspecific. More evidence is awaited. Bombus (Ps.) cornutus (Frison) cornutus (Frison, 1933:338 [Psithyrus]) pyramideus (Maa, 1948:19 [Psithyrus]) examined acutisquameus (Maa, 1948:21 [Psithyrus]) examined Klapperichi (Pittioni, 1949:273 [Psithyrus]) examined, not of Pittioni, 1949:266 (= B. picipes Richards) ?canus (Tkalcit, 1989:42 [Psithyrus]) Bombus (Ps.) expolitus Tkalcit expolitus (Tkalct, 1989:44 [Psithyrus]) examined Bombus (Ps.) turneri (Richards) turneri (Richards, 1929a:141 [Psithyrus]) examined ?monozonus (Friese, 1931:304 [Psithyrus]) not of Friese, 1909:674 (= B. lucorum (Linnaeus)) ?decoomani (Maa, 1948:26 [Psithyrus]) examined ?martensi (Tkalct, 1974b:314 [Psithyrus]) (provisional synonym) @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species. However, aside from differences in colour pattern, they are closely similar in morphology. Until more evidence to the contrary is available from critical studies of pat- terns of variation, I shall treat them as parts of a single variable species. Bombus (Ps.) tibetanus (Morawitz) tibetanus (Morawitz, 1886:202 [Apathus]) ?latefasciatus (Friese, 1931:304 [Psithyrus]) Bombus (Ps.) chinensis (Morawitz) chinensis (Morawitz, 1890[April 30]:352 [Apathus]) morawitzi (Friese, 1905:516 [Psithyrus]) not of Radoszkowski, 1876:101 (= B. morawitzi Rado- szkowski) hénei (Bischoff, 1936:26 [Psithyrus]) not of Bischoff, 1936:10 (= B. friseanus Skorikoy) CHECKLIST OF BUMBLE BEES Bombus (Ps.) novus (Frison) novus (Frison, 1933:340 [Psithyrus]) ?nepalensis (Tkalcti, 19746:318 [Psithyrus]) examined Bombus (Ps.) branickii (Radoszkowski) Branickii (Radoszkowski, 1893:241 [Psithyrus]) exam- ined chloronotus (Morawitz, 1894:6 [Apathus]) elisabethae (Reinig, 1940:231 [Psithyrus]) examined [branichi (Kim & Ito, 1987:32 [Psithyrus]) incorrect sub- sequent spelling] Bombus (Ps.) rupestris (Fabricius) rupeftris (Fabricius, 1793:320 [Apis]) Pyrencus (Lepeletier, 1832:375 [Psithyrus]) Interruptus (Lepeletier, 1832:381 [Psithyrus]) armeniacus (Reinig, 1970:77 [Psithyrus]) not of Radoszkowski, 1877b:202 (= B. armeniacus Radoszkowski) Bombus (Ps.) ferganicus (Radoszkowski) ferganicus (Radoszkowski, 1893:241 [Psithyrus]) exam- ined ochraceus (Morawitz, 1894:5 [Apathus]) indicus (Richards, 1929a:139) examined Bombus (Ps.) morawitzianus (Popov) morawitzianus (Popov, 1931:148,183 [Psithyrus]) exam- ined redikorzevi (Popov, 1931:160,181 [Psithyrus]) O NOMENCLATURE. Griitte (1937) regarded B. morawitzianus and B. redikorzevi as conspecific and, following the Principle of First Reviser (ICZN, 1985: Article 24), chose B. morawitzianus as the name for the species. Bombus (Ps.) campestris (Panzer) campestris (Panzer, 1801(74):11 [Apis]) Varius (Lepeletier, 1832:381 [Psithyrus]) flavus (Pérez, 1884:265 [Psithyrus]) flavo-thoracicus (Hoffer, 1889:49 [Psithyrus]) ?Susterai (Tkalcti, 1959:251 [Psithyrus]) examined, not of May, 1944:267 (= B. maxillosus Klug) (provisional synonym) ?susteraianus (Tkalcu, 1977:224 [Psithyrus]) replacement name for susterai Tkalcii, 1959:251 (provisional syno- nym) @ TAXONOMIC stTaTUS. Iam unaware of any rea- son (other than minor differences) why B. campestris and B. susteraianus should not be considered conspecific. 105 Bombus (Ps.) bellardii (Gribodo) Bellardti (Gribodo, 1892:108 [Psithyrus]) examined pieli (Maa, 1948:29 [Psithyrus]) examined, new synonym tajushanensis (Pittioni, 1949:277 [Psithyrus]) examined, not of Pittioni, 1949:244 (= B. kulingensis Cockerell), new synonym @ TAXONOMIC sTATUS. B. bellardii, B. pieli and B. tajushanensis are closely similar in morphology and I am unaware of any reason why these nominal taxa should not be considered conspecific. O NOMENCLATURE. For this species, the oldest available name is B. bellardii, which becomes the valid name. The only subsequent publications using the name B. pieli of which I am aware are by Maa (1948), Sakagami (1972), Tkalciit (1987) and Williams (1991), so this change of valid name is not a serious disruption of common usage. Bombus (Ps.) norvegicus (Sparre-Schneider)® norvegicus (Sparre-Schneider, 1918:40 [Psithyrus]) not of Friese, 1911:571 (= B. monticola Smith) transhaicalicus (Popov, 1927:269 [Psithyrus]) O NOMENCLATURE. With Psithyrus regarded as being a subgenus of the genus Bombus (Williams, 1991, 1995), P. norvegicus Sparre-Schneider (1918) becomes a junior secondary homonym in Bombus of B. lapponicus var. norvegicus Friese (1911) (deemed subspecific, see ICZN, 1985: Article 45g(ii)), and therefore the name P. norvegicus Sparre-Schneider is invalid (ICZN, 1985: Article 57c). For this species, the oldest available name of which I am aware is P. norvegicus var. transbaicalicus Popov, 1927 (deemed to be subspecific, see ICZN, 1985: Article 45g(ii)), so B. transbaicalicus would become the valid name. © APPLICATION TO ICZN. Although B. trans- baicalicus is the oldest available name for this species, the name B. norvegicus has been in com- mon use for the species since 1947 (e.g. Faester & Hammer, 1970; Delmas, 1976; Ito & Tadauchi, 1981; Pekkarinen ef al., 1981; Reinig, 1981; Loken & Framstad, 1983; Rasmont, 1983; Loken, 1984; Ito, 1985; Pekkarinen & Teras, 1993; Rasmont ef al., 1995). It is suggested that, in the interests of stabil- ity, an application be made to ICZN to use its Plenary Power to suppress the senior homonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). However, the consequence of this action would be that norvegicus Friese would no longer be available for a subspecies of B. monticola. Bombus (Ps.) fernaldae (Franklin) fernalde (Franklin, 1911:164 [Psithyrus]) examined 106 Bombus (Ps.) flavidus Eversmann flavidus Eversmann, 1852:131 lissonurus (Thomson, 1872:49 [Apathus]) @ TAXONOMIC STATUS. Rasmont (1988) reports that the Pyrenean population of B. flavidus is morphometrically distinct from the disjunct Scan- dinavian population (comparable distinctions are not known within its close relatives B. norvegicus and B. sylvestris, which share these areas of distribtuion). Nevertheless he continues to treat them as conspecific and I shall follow this, at least until further evidence in support of two separate species is available. Bombus (Ps.) skorikovi (Popov) skorikovi (Popov, 1927:267 [Psithyrus]) examined ?gansuensis (Popov, 1931:202 [Psithyrus]) ?kuant (Tkalcu, 1961b:362 [Psithyrus]) Bombus (Ps.) quadricolor (Lepeletier) Quadricolor (Lepeletier, 1832:376 [Psithyrus]) globosus (Eversmann, 1852:126 [Psithyrus]) meridionalis (Richards, 1928b:351 [Psithyrus]) not of Dalla Torre, 1879:13 (= B. hortorum (Linnaeus)) Bombus (Ps.) sylvestris (Lepeletier) Sylvestris (Lepeletier, 1832:377 [Psithyrus]) Brasiliensis (Smith, 1854:385 [Apathus]) examined, not of Lepeletier, 1836:470 (= B. brasiliensis Lepeletier) citrinus (Schmiedeknecht, 1883[see Baker, 1996c:297]:23[407] [Psithyrus]) not of Smith, 1854:385 (= B. citrinus (Smith)) [silvestris (Dalla Torre, 1896:571 [Psithyrus]) incorrect subsequent spelling] e00e000ee0e er Pats eecsece: * ee se 00% oe one f $7 Subgenus LAESOBOMBUS Kriiger Bombus (Laesobombus) Kriiger, 1920:350, type-species Bombus laesus Morawitz by monotypy Agrobombus (Laesobombus) Skorikov, 1922b:20, type- species Bombus laesus Morawitz by monotypy Agribombus (Laesibombus) Skorikoy, 1938a:145, unjusti- fied emendation P.H. WILLIAMS Bombus (Ls.) laesus Morawitz laesus Morawitz in Fedtschenko, 1875:3 Mocsaryi Kriechbaumer, 1877:253 ?maculidorsis (Skorikoy, 1922b:23 [Agrobombus]) not infrasubspecific after Panfilov, 1956:1328 ?tianschanicus Panfilov, 1956:1327 (provisional synonym) ferrugifer Reinig, 1971:158 @ TAXONOMIC STATUS. Panfilov (1956) regarded B. laesus, B. mocsaryi, B. maculidorsis and B. tian- schanicus as separate species, differing particularly in: (1) the colour of the pubescence on the thoracic dor- sum; (2) the number of large punctures on the clypeus; (3) the strength of the median keel on gastral sternum VI; and (4) the length of the hair of the dorsum. However, from the material I have examined (collec- tions in London, Beijing), these character states do not appear to be either discreet or strongly associated. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Subgenus ORIENTALIBOMBUS Richards Bombus (Orientalibombus) Richards, 1929c:378, type- species Bombus orientalis Smith (= Bombus haemorrhoidalis Smith) by original designation Bombus (Orientalobombus) Kruseman, 1952:102, unjus- tified emendation Bombus (Or.) funerarius Smith funerarius Smith, 1852b:47, examined priscus (Frison, 1935:349 [Bremus]) birmanus (Tkalct, 1989:47 [Orientalibombus]) examined Bombus (Or.) braccatus Friese braccatus Friese, 1905:512, examined metcalfi (Frison, 1935:357 [Bremus]) examined Bombus (Or.) haemorrhoidalis Smith heemorrhoidalis Smith, 1852a:43 orientalis Smith, 1854:402, examined assamensis Bingham, 1897:550, examined CHECKLIST OF BUMBLE BEES montivolans Richards, 1929c:382, examined semialbopleuralis (Tkalcit, 1974b:322 [Orientalibombus]) cinnameus (Tkalcii, 1989:47 [Orientalibombus]) examined @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species, most recently in the case of B. montivolans [Burma to southern China] (e.g. Tkalcii, 1968b, 1989). However, aside from differ- ences in colour pattern, they are all closely similar in morphology with a range of variation (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Subgenus EXILOBOMBUS Skorikov Mucidobombus (Exilobombus) Skorikov, 1922a:150, type- species Mucidobombus exil Skorikoy (cited as exiln.) (= Bombus exil (Skorikoy)) by monotypy Megabombus (Exilnobombus) Milliron, 1973a:81, unjus- tified emendation Bombus (Ex.) exil (Skorikov) exiln. nov. (Skorikovy, 1922a:150 |[Mucidobombus}) {not a replacement name] [exul (Skorikov, 1931:216 [Mucidobombus}) incorrect sub- sequent spelling] exil (Milliron, 1961:56 [Megabombus]) justified emenda- tion [exilis Richards, 1968:254, incorrect subsequent spelling] exul (Tkalcti, 1974a:42 [Megabombus]) unjustified emen- dation Subgenus THORACOBOMBUS Dalla Torre Bombus (Thoracobombus) Dalla Torre, 1880:40, type- 107 species Apis sylvarum Linnaeus (= Bombus sylvarum (Linnaeus)) by subsequent designation of Sandhouse, 1943:604 Bombus (Chromobombus) Dalla Torre, 1880:40, type-spe- cies Apis muscorum Linnaeus (= Bombus muscorum (Linnaeus)) by subsequent designation of Sandhouse, 1943:538 Bombus (Agrobombus) Vogt, 1911:52, type-species Apis agrorum Fabricius (=Bombus pascuorum (Scopoli)) by subsequent designation of Sandhouse, 1943:523 [Agrabombus Skorikov, 1914a:119, incorrect subsequent spelling] Bombus (Ruderariobombus) Kriiger, 1920:350, type-spe- cies Apis ruderaria Miiller (= Bombus ruderarius (Miiller)) by subsequent designation of Yarrow, 197 1:27 Agrobombus (Adventoribombus) Skorikoy, 1922a:150, type-speciesA grabombus adventor Skorikov (=Bombus filchnerae Vogt) by subsequent designation of Sandhouse, 1943:522, new synonym [Agrobombus (Adventoriobombus) Skorikoy, 1931:218, incorrect subsequent spelling] Agribombus Skorikovy, 1938a:145, unjustified emendation [Bombus (Thoraocbombus) Esmaili & Rastegar, 1974:52, incorrect subsequent spelling] [Bombus (Thoracibombus) Schwarz et al., 1996:197, in- correct subsequent spelling] @ TAXONOMIC STATUS. Richards (1968) treated Thoracobombus and Adventoribombus as separate subgenera, although he questioned whether they should be kept separate. I have followed Tkalcti (1974a) in treating B. adventor (=B. filchnerae) as part of a single subgenus Thoracobombus. Bombus (Th.) filchnerae Vogt Filchnerae Vogt, 1908:100, examined adventor (Skorikov, 1914a:119 [Agrabombus}) lii Tkalcit, 1961b:355 Bombus (Th.) muscorum (Linnaeus) Mujcorum (Linnaeus, 1758:579 [Apis]) examined pallidus Evans, 1901:47, not of Cresson, 1863:92 (= B. pensylvanicus (DeGeer)) [fulvofasciatus Friese, 1905:520, infrasubspecific] laevis Vogt, 1909:63 ?nigripes Pérez, 1909:158, not of Haliday in Curtis ef al., 1837:321 (= B. dahlbomii Guérin-Méneville) ?pereziellus (Skorikoy, 1922a:150 [Agrobombus]) replace- ment name for nigripes Pérez, 1909:158 ?bannitus (Skorikov in Popov, 1930:98 [Agrobombus]) ?liepetterseni Loken, 1973:152 celticus Yarrow, 1978:15, replacement name for pallidus Evans, 1901:47 @ TAXONOMIC STATUS. B. bannitus (= B. smith- ianus of authors, a misidentification (=B. pascuorum)) has been regarded as a separate species by some authors (e.g. Richards, 1935; Tkalct, 1987; Rasmont & Adamski, 1995) on the basis of its semi-melanic colour pattern and more coarsely sculptured surface of 108 gastral terga [V-V. However, Loken (1973: fig. 81) found no difference between these taxa in a morphometric study (other authors reporting no clear morphological differences include Richards, 1935; Alford, 1975; Pekkarinen, 1979; Rasmont, 1982; Baker, 1996a). Furthermore, I have collected many specimens with a range of intermediate colour patterns on the Isle of Skye in western Scotland. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. B. pereziellus has also been regarded as a separate species by Rasmont & Adamski (1995), because of its dark colour pattern (even darker than B. bannitus, B. pereziellus has the thoracic dorsum black rather than red-brown, and has more black hairs on gastral tergum II, whereas these black hairs tend to be more frequent on tergum I for B. bannitus) and because it is endemic to the island of Corsica. Morphologically it was con- sidered by Rasmont (1982) to show no perceptible differences from B. muscorum or B. bannitus. Further- more, a male with a colour pattern apparently intermediate between B. muscorum and B. pereziellus is mentioned by Delmas (1976:271). Depending on the species concept embraced, some differences might be expected for a peripheral population such as this even if it were conspecific and I shall treat them as parts of a single variable species. Further evidence is awaited. O NOMENCLATURE. Richards (1935, 1968), Yarrow (1968) and Léken (1973) recognised that none of the admissable syntypes in the Linnean collection agreed with the traditional interpretation of B. muscorum, which is very rare in the parts of Sweden where Linnaeus collected (Richards, 1935; Loken, 1973; Day, 1979), but took no action. When Day (1979) came to fix the application of the name, he had no reason to believe that Linnaeus had not described his A. muscorum from the syntype specimen that was subsequently described as lectotype (= B. humilis Illiger). To reaffirm the traditional usage of B. muscorum, a case was made to ICZN by Loken et al. (1994). This sought an Opinion from ICZN (ICZN, 1996) that set aside by use of its Plenary Power (ICZN, 1985: Arti- cles 78b, 79) the lectotype designation forA. muscorum by Day from application of the Code (ICZN, 1985) and then designated a neotype (ICZN, 1996: 64) to conserve the traditional usage of the name for even the narrowest concept of the taxon (ICZN, 1985: Article 75). Bombus (Th.) anachoreta (Skorikov) anachoreta (Skorikov, 1914a:121 [Agrobombus]) P.H. WILLIAMS Bombus (Th.) opulentus Smith opulentus Smith, 1861:153, examined Bombus (Th.) zonatus Smith zonatus Smith, 1854:389 Bombus (Th.) humilis MligerS fulvefcens (Schrank, 1802:367 [Apis]) humilis Wliger, 1806:171 ?tristis Seidl, 1837:69 ?variabilis Schmiedeknecht, 1878:424, not of Cresson, 1872:284 (= B. variabilis (Cresson)) ?subbaicalensis Vogt, 1911:42,54 OQ NOMENCLATURE. When Day (1979) came to fix the application of A. muscorum Linnaeus (see the comments on B. muscorum), he had no reason to believe that Linnaeus had not described this taxon from the syntype specimen that was subsequently described as lectotype (= B. humilis Mlliger). This action brought B. humilis Mliger into subjective junior synonymy with B. muscorum (Linnaeus). To reaffirm the traditional usage of B. muscorum and B. humilis, a case was made to ICZN by Loken et al. (1994). This sought an Opinion from ICZN (ICZN, 1996) that set aside by use of its Plenary Power (ICZN, 1985: Articles 78b, 79) the lectotype designation forA. muscorum by Day from application of the Code (ICZN, 1985) and then designated a neotype (ICZN, 1996: 64) to conserve the traditional usage of B. muscorum and B. humilis ((CZN, 1985: Article 75). However, Warncke (1986) recognised B. fulvescens (Schrank) as questionably conspecific with B. humilis. I have seen no type specimens, but the description is consistent with this interpretation. B. fulvescens is therefore likely to be the oldest available name for this species. € APPLICATION TO ICZN. Although B. fulvescens may be the oldest available name for the present interpretation of this species, the name B. humilis has been in common use for the species since 1947 (e.g. case and references in Lgken et al., 1994). In contrast, I know of no publications using the name B. fulvescens (Schrank) since 1947. Warncke (1986:98) followed the listing of this name with ‘Art. 23b’, which is a reference to purpose of the Principle of Priority (ICZN, 1985). I agree that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). Bombus (Th.) deuteronymus Schulz senilis Smith, 1879:131, examined, not of Fabricius, 1775:382 (= B. pascuorum (Scopoli)) deuteronymus Schulz, 1906:267, replacement name for CHECKLIST OF BUMBLE BEES senilis Smith, 1879:131 velox (Skorikoy, 1914a:120 [Agrobombus}]) [superequester (Skorikov, 1914c:405 [Agrobombus]) infrasubspecific ] superequester (Skorikov, 1925:116 [Agrobombus}) bureschi Pittioni, 1939b:1, examined Bombus (Th.) schrencki Morawitz Schrencki Morawitz, 1881:123 Schrencki Morawitz, 1881:250, redescribed konakovi Panfiloy, 1956:1330 ?Bombus (Th.) honshuensis (Tkalcii) honshuensis (Tkalcti, 1968a:47 [Megabombus]) @ TAXONOMIC stTATUS. B. honshuensis and B. schrencki have allopatric distributions in northern Ja- pan (Sakagami & Ishikawa, 1969; Ito & Munakata, 1979: fig. 6; Ito, 1993), with B. honshuensis being possibly a disjunct peripheral population of B. schrencki. The two taxa are closely similar, and yet despite some variation in morphology, apparently con- sistent differences have been described (Tkalcii, 1968a; Sakagami & Ishikawa, 1972). Nonetheless, some dif- ferences might be expected even if they were conspecific, depending on the species concept ac- cepted (see the comments on B. ruderatus), so further evidence is awaited. Bombus (Th.) impetuosus Smith impetuosus Smith, 1871:249, examined Potanini Morawitz, 1890:350, new synonym yuennanensis Bischoff, 1936:14, examined combai Tkalcti, 1961b:357, new synonym @ TAXONOMIC STATUS. The white-banded B. potanini is morphologically closely similar to the yellow-banded B. impetuosus. Some individuals from Sichuan are intermediate in colour pattern in that they have the pale bands of the thorax and gastral tergum I white, and the pale band of tergum II yellow. There is considerable variation in the male gonostylus, but this variation appears to overlap between the the colour forms and I shall treat them as parts of a single variable species. S.-f. Wang and J. Yao (in litt.) also believe that the two taxa may be conspecific. Further evidence is awaited. Bombus (Th.) remotus (Tkalcii) remotus (Tkalcti, 1968a:45 [Megabombus]) examined Bombus (Th.) pseudobaicalensis Vogt Pseudobaicalensis Vogt, 1911:43,53 gilvus (Skorikov, 1925:117 [Agrobombus}) 109 Bombus (Th.) hedini Bischoff unicolor Friese, 1905:514, examined, not of Kriechbaumer, 1870:159 (= B. maxillosus Klug) hedini Bischoff, 1936:15 Bombus (Th.) ruderarius (Miiller) ruderaria (Miller, 1776:165 [Apis]) Derhamella (Kirby, 1802:363 [Apis]) examined montanus Lepeletier, 1836:463 simulatilis Radoszkowski, 1888:317, examined Bombus (Th.) inexspectatus Tkalcit lutescens Kriiger, 1939:105, not of Pérez, 1890:154 (= B. flavidus Eversmann) inexspectatus (Tkalct, 1963:187 [Agrobombus}) [inexpectatus (Reinig, 1981:161 [Megabombus}]) incorrect subsequent spelling] COMMENT. On the grounds of its peculiar morphol- ogy, B. inexspectatus has been suggested to be an obligate social parasite in colonies of other Bombus species, with B. ruderarius being the most likely host (Yarrow, 1970). As yet, there is no direct evidence for this behaviour (Rasmont, 1988). See the comments on the subgenus Psithyrus and on B. hyperboreus. Bombus (Th.) sylvarum (Linnaeus) /ylvarum (Linnaeus, 1761:425 [Apis]) examined Daghestanicus Radoszkowski, 1877a:vii Dagestanicus Radoszkowski, 1877b:211, redescribed Bombus (Th.) veteranus (Fabricius) veterana (Fabricius, 1793:324 [Apis]) arenicola Thomson, 1872:31 Bombus (Th.) mlokosievitzii Radoszkowski Mlokosievitzii Radoszkowski, 1877a:viii Mlokassewiczi Radoszkowski, 1877b:212, redescribed pérezi Vogt, 1911:55, not of Schulthess-Rechberg, 1886:275 (= B. perezi (Schulthess-Rechberg)) vogtiellus (Tkalcu, 1977:224 [Megabombus]) replacement name for perezi Vogt, 1911:55 [mlokossowiczi (Reinig, 1981:161 [Megabombus]) incor- rect subsequent spelling] O NOMENCLATURE. There are particularly many incorrect subsequent spellings of B. mlokosievitzii. Bombus (Th.) pascuorum (Scopoli) Pafcuorum (Scopoli, 1763:306 [Apis]) Jenilis (Fabricius, 1775:382 [Apis]) agrorum (Fabricius, 1787:301 [Apis]) not of Schrank, 1781:397 (= B. mesomelas Gerstaecker) thoracicus Spinola, 1806:30 110 arcticus Dahlbom, 1832:50, not of Quenzel in Acerbi, 1802:253 (= B. hyperboreus Schonherr) cognatus Stephens, 1846:17, examined smithianus White, 1851:158 @ TAXONOMIC STATUS. Warncke (1986) listed B. cognatus as a synonym of B. muscorum, possibly following Stephens (1846), who wrote of B. cognatus: ‘Closely allied to Bo. Muscorum, of which the exam- ples I possess may be immature specimens’. Pagliano (1995) listed B. cognatus as a species separate from both B. muscorum and B. pascuorum, but without any explanation. Saunders (1896:366—367) wrote ‘I have re-exam- ined the type of cognatus, Steph., . . . F. Smith placed it in the British Museum collection. ... Saunders considered this specimen to be conspecific with B. agrorum (Fabricius), continuing: ‘It is certainly not the species known on the Continent as cognatus’. A female in the NHM collection bears the following labels: (1) a red-edged printed ‘Type’; (2) ‘cognatus.’ in handwriting identical to that of F. Smith; (3) ‘= agrorum / I.H.H.Y.’ in handwriting identical to that of I. Yarrow; (4) ‘B.M. Type / HYM. / 17B.1163’. I have examined this specimen and am unaware of any reason why it should not be considered the type of B. cogna- tus and (ignoring minor differences in colour pattern) conspecific with B. pascuorum. OQ NOMENCLATURE. L@ken (1973) listed B. cogna- tus Stephens, 1846, as anomen nudum, citing Sherborn (1925). However, the reference by Sherborn is to an earlier paper by Stephens (1829), so this does not affect the use of the name B. cognatus Stephens, 1846. Subgenus TRICORNIBOMBUS Skorikov Agrobombus (Tricornibombus ) Skorikoy, 1922a:151, type- species Bombus tricornis Radoszkowski by monotypy Bombus (Tricornibombus) Tkalcti, 1960:70 Bombus (Tr.) tricornis Radoszkowski tricornis Radoszkowski, 1888:319, examined Bombus (Tr.) atripes Smith atripes Smith, 1852a:44, examined P.H. WILLIAMS Bombus (Tr.) imitator Pittioni imitator Pittioni, 1949:251, examined flavescens Pittioni, 1949:254, not of Smith, 1852a:45 (=B. flavescens Smith) Subgenus FERVIDOBOMBUS Skorikov Fervidobombus Skorikov, 1922a:153, type-species Apis fervida Fabricius (= Bombus fervidus (Fabricius)) by subsequent designation of Frison, 1927:69 Bombus (Fervidobombus) Franklin, 1954:47 Bombus (Digressobombus ) Laverty etal., 1984:1051, type- species Megabombus digressus Milliron (= Bombus digressus (Milliron)) by original designation @ TAXONOMIC STATUS. The subgenus Digresso- bombus was described subsequent to Richards (1968). I have treated Digressobombus as part of a single subgenus Fervidobombus (Williams, 1995), as has Labougle (1990). While this study found no evidence for monophyly of the combined group, I believe that this is more likely (unpublished data) than monophyly of the subgenus Fervidobombus excluding Digresso- bombus. COMMENT. This is the only early-diverging and large subgenus of bumble bees to occur in the New World other than the subgenus Psithyrus. Although it makes up only a small part of the fauna of America north of Mexico, it makes up most of the low- to mid-altitude bumblebee fauna of Central and SouthAmerica. It also includes the only species of bumble bees genuinely occurring in tropical lowland wet forest (e.g. Moure & Sakagami, 1962; Milliron, 1973a; Cameron & Whitfield, 1996). The species with more temperate distributions appear to occupy similar habitats and show similar flower-depth preferences to species of subgenera such as Thoracobombus and Megabombus in the Old World. Bombus (Fy.) fervidus (Fabricius) feruida (Fabricius, 1798:274 [Apis]) ?Californicus Smith, 1854:400, examined Dumoucheli Radoszkowski, 1884:78 sonome Howard, 1902:pl. II @ TAXONOMIC STATUS. B. fervidus and B. CHECKLIST OF BUMBLE BEES californicus have been regarded both as conspecific (e.g. Milliron, 1973a; Labougle, 1990) and as separate species (e.g. Franklin, 1913; Stephen, 1957; Thorp et al., 1983: Poole, 1996). Both Franklin (1913:239) and Stephen (1957) also considered the possibility that they are conspecific as quite reasonable. Many specimens from the north west of North America show a reduction in the extent of the yellow bands on the scutellum and gastral terga I-III and appear to be intermediate or recombinant individuals. Indeed, Stephen’s (1957:32) figure 2 shows several patterns that could represent a continuum in variation between the two forms. Thorp ef al. (1983) found no intermediate females in California, although some of the males of B. californicus were said to approach the pattern of B. fervidus. In view of the existence of apparent intermediates between these nominal taxa in at least part of their range, they are treated here as likely to be conspecific. More evidence is awaited. OQ NOMENCLATURE. Apis feruida is the original spelling in Fabricius (1798). The orthography of this publication employs ‘u’ in place of “v’ widely, a com- mon practice of the period. This convention has since changed and subsequent authors have consistently used ‘vy’ for B. fervidus. In fact, whatever the interpretation of the Code, pragmatically it matters little which spelling offervidus is used unless either of the spellings were to be pub- lished as the name of another taxon in Bombus. See the comments on the spelling of B. pensylvanicus. Bombus (Fy.) pensylvanicus (DeGeer) penfylvanica (DeGeer, 1773:575 [Apis]) americanorum Fabricius, 1804:346 ?sonorus Say, 1837:413 pallidus Cresson, 1863:92 Pensylvanicus Cresson, 1863:94 flavodorsalis Franklin, 1913:409 pennsylvanicus Hurd, 1979:2204, unjustified emendation @ TAXONOMIC sTATUS. B. pensylvanicus and B. sonorus have been regarded both as conspecific (e.g. Milliron, 1973a; Labougle ef al., 1985; Labougle, 1990; Poole, 1996) and as separate species (e.g. Franklin, 1913 [but see p. 239]; Stephen, 1957; Thorp et al., 1983; S. Cameron in litt.). From the few males from the United States (not Mexico) that I have examined in detail, there appear to be subtle differences in the male genitalia (e.g. in the shape of the penis valve head). However, Labougle (1990) reports that the two ‘forms are geographically intermixed in México, and chromatically intermediate specimens occur, mainly in northeastern México and southwestern Texas’. He went on to say that ‘In fact, it is sometimes difficult to place a Mexican specimen in either subspecies because there are specimens with the 111 coloration of the scutellum and the punctation of the clypeus intermediate between the two taxa. Average differences of certain proportions are found . . . but do not differentiate all specimens’. G. Chavarria (pers. com.) also believes that intermediate specimens occur in Mexico and that they are conspecific. Taking an extreme viewpoint, it is even possible to see the ‘typi- cal’ B. sonorus colour pattern as intermediate between B. pensylvanicus (in the strict sense) and the extreme pale form that has the thoracic dorsum and gastral tergum I entirely yellow (flavodorsalis, see Thorp et al., 1983: fig. 137b). In view of the existence of apparent intermediates between these nominal taxa in at least part of their range, they are treated here as likely to be conspecific. More evidence is awaited. O NOMENCLATURE. Apis penfylvanica is the origi- nal spelling in DeGeer (1773). The orthography of this publication employs ‘Pin place of ‘s’ widely, a com- mon practice of the period. This convention has since changed and subsequent authors (e.g. Cresson, 1863) have consistently used ‘s’ for B. pensylvanicus. Technically, according to the Code (ICZN, 1985: Article 32b), pensylvanicus with just two ‘n’s is the correct original spelling, to be preserved unaltered unless it is demonstrably incorrect under Article 32c. Article 32c(ii) states that clear evidence of an inadvert- ent error is only admissable if it lies within the original publication, without recourse to any external source of information (DeGeer, 1773, spelled Penjylvanie and penfylvanica consistently in this way). Any intentional change to that spelling in a subsequent publication is an unjustified emendation under Article 33b(iii). In fact, whatever the interpretation of the Code, pragmatically it matters litthe which spelling of pensylvanicus is used unless either of the spellings were to be published as the name for another taxon in Bombus. No doubt many will prefer to use B. pennsylvanicus, although the name does appear as B. pensylvanicus in the recent checklist by Poole (1996) (and by analogy, the similar spelling of Vespula pensylvanica (Saussure) has been accepted, e.g. by Akre et al., 1980; Edwards, 1980). COMMENT. This species was deliberately introduced into the Philippines, but is not known to have persisted (Frison, 1925b). Bombus (Fy.) excellens Smith excellens Smith, 1879:133, examined Bombus (Fy.) dahlbomii Guérin-Méneville Dahlbomii Guérin-Méneville, [1835, see Cowan, NO TE2 epi nigripes Haliday in Curtis et al., 1836:321 112 O NOMENCLATURE. Cowan (1971), considering Guérin-Méneville’s insect volume, states that ‘it is quite certain that valid publication [of the Insectes text] under the International Code of Nomenclature did not take place until August or September 1844.’ However, he lists plate 75, on which B. dahlbomii appears as figure 3 together with a legend containing the name, as having been published in livraison 39 in June 1835. This meets the criteria for valid publication (ICZN, 1985: Article 8). Therefore B. dahlbomii ts the oldest available name for this species. Bombus (Fyv.) morio (Swederus) morio (Swederus, 1787:283 [Apis]) examined velutinus Mliger, 1806:175 violaceus Lepeletier, 1836:473 carbonarius Handlirsch, 1888:241, not of Menge, 1856:27 [fossil] Kohli Cockerell, 1906:75, replacement name for carbonarius Handlirsch, 1888:241 Bombus (Fy.) diligens Smith diligens Smith, 1861:154, examined dolichocephalus Handlirsch, 1888:244 Bombus (Fy.) opifex Smith opifex Smith, 1879:133, examined Bombus (Fy.) rubriventris Lepeletier rubriventris Lepeletier, 1836:472, examined @ TAXONOMIC sTATUS. B. rubriventris is known from a single female specimen from ‘St. Domingue’ (2= Sao Domingos, Goids) (Milliron, 1973a). This specimen has dark brown wings and the pubescence is black, except that most of the hairs of the thorax are grey-tipped, and the hairs of gastral terga II-IV are bright ‘coppery’ red. This colour pattern resembles the Andean B. excellens, although the pubescence of B. rubriventris is much shorter and more even; the oculo-malar area is nearly square rather than nearly twice as long as the basal breadth of mandible; and tergum VI is raised subapically. Franklin (1913) had not seen B. rubriventris but suggested that it was probably a ‘freak specimen’ of B. carolinus (a misidentification, = B. excellens). Milliron (1973a) had examined B. rubriventris and considered the morphological char- acters to be very much like those of B. bellicosus. However, B. rubriventris can be distinguished by the much finer punctures in the centre of the clypeus and by an absence of a median ridge on tergum VI. I consider B. rubriventris to be more similar in these characters to B. opifex, although it can be distin- P.H. WILLIAMS guished from that species by a pair of characteristi- cally slightly recessed bands of fine punctures extending anteriorly from the ocello-ocular areas and by a shallow median groove in the subapically raised area of tergum VI. The colour pattern is very distinctive among non- Andean bumble bees in South America and does not appear to be the result of abnormal colour develop- ment. The specimen has had the gaster glued back into place, although the characters of both the head and gaster appear to be distinctive, so there is no reason to believe that the specimen is a composite and not genuine. COMMENT. Milliron (1973a) researched the history of this specimen and believed that it may have been collected as early as 1800. He concluded that it was probably a highland species from southeastern Brazil and that it may now be extinct. If so, and accepting that it is very difficult to establish the absence of a species, this would be one of the few known cases of complete extinction of an insect species. Bombus (Fyv.) bellicosus Smith thoracicus Sichel, 1862:121, not of Spinola, 1806:30 (=B. pascuorum (Scopoli)) bellicosus Smith, 1879:131, examined Emiliae Dalla Torre, 1890:139, replacement name for thoracicus Sichel, 1862:121 Bombus (Fy.) pullatus Franklin pullatus Franklin, 1913:122 Bombus (Fy.) transversalis (Olivier) tran/verfalis (Olivier, 1789:65 [Apis]) Cajennenjis (Fabricius, 1798:273 [Apis]) incarum Franklin, 1913:131 Bombus (Fy.) atratus Franklin® azurea (Christ, 1791:129 [Apis]) ?atratus Franklin, 1913:118, not of Friese, 1911:572 (=B. mucidus Gerstaecker) (provisional synonym) ?niger Franklin, 1913:120, examined (provisional syno- nym) ?nigriventris Friese, 1913:87 (provisional synonym) @ TAXONOMIC stTaTUS. Atleast four species of the subgenus Fervidobombus from Central and South America have many individuals for which the pubes- cence is almost entirely black. The genitalia of the males are quite distinctive, but association of the conspecific females with these males has caused prob- lems. In the original description of B. niger, Franklin stated that ‘atratus is possibly the male of niger (p. CHECKLIST OF BUMBLE BEES 121), whereas in the original description of B. atratus he stated both that ‘Niger may represent the females of this species’ (p. 118) and that “This may be the true male of kohli’ (p. 119). B. niger was described from a syntype series of four queens and four workers, of which one queen in the Smithsonian collection carries, amongst others, a red label ‘LECTOTYPE / Bombus / niger Franklin/H.E.Milliron ‘59’ and a label “Boquete / Chiriqui’. This Central American locality was men- tioned by Franklin, but is outside the known distribution of the species (Milliron, 1973a) to which the specimen belongs. In my opinion, this lectotype of B. niger is not conspecific with B. pullatus (contrary to the sugges- tion by Labougle, 1990, see also Milliron, 1962) but rather is conspecific with B. atratus Franklin. Another possibility is that this variable species is the Apis azurea of Christ (1791). I know of no type specimens and the type locality was said to be inAfrica (Ist in Afrika am Vorgebiirg der guten Hofnung zu Haus’). The description and figure of the colour pat- tern do not agree with any African bees that Ihave been able to trace, but do resemble closely the yellow- banded individuals of the SouthAmerican B. niger, the Central American B. medius Cresson, and the South American B. transversalis (Olivier) (although for the last named species the yellow bands on the thorax are usually broader). Among the specimens to hand, the wings do appear slightly more “Schwarzblaue’ for B. niger, as described for A. azurea, although these grounds seem slim justification from which to estab- lish the application of a name. O NOMENCLATURE. B. azureus is possibly the old- est available name for this species. Milliron (1962), without mention of the name B. azureus, first regarded B. atratus and B. niger as conspecific and, following the Principle of First Re- viser (ICZN, 1985: Article 24), chose B. atratus as the valid name for the species. Unfortunately, B. atratus Franklin, 1913, is a junior primary homonym of B. mucidus var. atratus Friese, 1911 (deemed to be subspecific, see ICZN, 1985: Article 45g(ii)), therefore the name B. atratus Franklin is invalid (ICZN, 1985: Article 57b). © APPLICATION TOICZN. The name B. azureus has not been used since the original publication. The name B. atratus has been used for this species since 1947 (e.g. Moure & Sakagami, 1962; Sakagami & Zucchi, 1965; Sakagami ef al., 1967; Milliron, 1971, 1973a; Sakagami, 1976; Ito, 1985; Labougle, 1990; Varela, 1992; Silveira & Cure, 1993). It is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress both the unused senior synonym (ICZN, 1985: Article 79) and the senior homonym. This would achieve both an unambiguous, valid name for this species (see the comments on B. muscorum) and also 113 help to protect the validity of the names B. medius and B. transversalis from future change. However, the consequence of this action would be that atratus Friese would no longer be available for a subspecies of B. mucidus. Bombus (Fv.) digressus (Milliron) digressus (Milliron, 1962:730 [Megabombus]) examined Bombus (Fy.) brasiliensis Lepeletier brasiliensis Lepeletier, 1836:470, examined Bombus (Fy.) steindachneri Handlirsch Steindachneri Handlirsch, 1888:239 @ TAXONOMIC sTATUS. B. medius andB. steindac- hneri have been regarded both as separate species (Milliron, 1973a; Labougle, 1990) and as conspecific (G. Chavarria, pers. com.). Labougle (1990) reports that ‘Although the chro- matic differences between B. medius and B. steindachneri are conspicuous, the male genitalia are extremely similar’. Labougle listed four character dif- ferences from the male genitalia and I can confirm two of these: (1) that the head of the penis valve of B. steindachneri has fewer fine teeth or serrations; and (2) that the interior process of the volsella (misinter- preted as the preapical tooth of the “gonostylus’; for discussion of homologies see Williams, 1991) of B. steindacheri is narrower. However, I have examined only a few males and these characters might be ex- pected to vary among other individuals. Labougle (1990) continued: ‘The lack of chromatic and morpho- logical intermediates supports the idea of two different species’. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as two separate species. Bombus (Fy.) medius Cresson medius Cresson, 1863:97 Bombus (Fy.) weisi Friese laboriosus Smith, 1861:153, examined, not of Fabricius, 1804:352 (= Emphoropsis laboriosus (Fabricius)) weisi Friese, 1903:253, examined nigrodorsalis Franklin, 1907:90 O NOMENCLATURE. The lectotype female of B. weisi by designation of Milliron (1960:98) was recog- nised as conspecific with B. nigrodorsalis by Labougle (1990) (Ihave examined the lectotype of B. weisi at the MNHU, Berlin, and agree with Labougle). He then 114 used B. weisi (the oldest available name) as the valid name for this species. However, a case could be made in favour of the use of either name. For Labougle’s (1990) use of this previously unused senior synonym to be considered by ICZN as a prima facie case of upsetting the use of a long-accepted name in its accustomed meaning (ICZN, 1985: Article 23b), the name B. weisi should not have been used in this sense in the preceding fifty years; and at least five authors should have used the junior name, B. nigrodorsalis, in at least ten publications during the same period (ICZN, 1985: Article 79c). As far as 1am aware, no other admissable publications have used B. weisi (Williams, 1995, disclaimed any nomenclatural action in a list of names for material examined), al- though publications using the junior name B. nigrodorsalis Franklin for this species since 1947 are more common, including Milliron (1961, 1962, 1971, 1973a), Laverty et al. (1984), Labougle et al. (1985), Williams (19855) andAsperen de Boer (1992b). Other such references may exist, therefore this may be seen as a borderline case, requiring an application to be made to ICZN to use its Plenary Power if suppression of the unused senior synonym, B. weisi, is required (see the comments on B. muscorum). On the other hand, a change of valid name from B. nigrodorsalis to B. weisi does not appear to be a serious disruption of common usage, so there is no obvious need for action to retain B. nigrodorsalis and I have continued to use B. weisi. Bombus (Fy.) trinominatus Dalla Torre modestus Smith, 1861:153, examined, not of Eversmann, 1852:134 (= B. modestus Eversmann) trinominatus DallaTorre, 1890:139, replacement name for modestus Smith, 1861:153 xelajuensis Asperen de Boer, 1992b:162, examined (pro- visional synonym) @ TAXONOMIC STATUS. The description of B. xelajuensis shows that this nominal taxon, known from a single location, diverges only slightly in col- our pattern and morphology from the otherwise restricted and uncommon mountain species B. trinominatus. Therefore it seems most likely to be conspecific with B. trinominatus, with a slightly dif- ferent colour pattern. However, the information available at present is not conclusive, and it remains possible that it represents a separate species, and further evidence is awaited. Bombus (Fy.) mexicanus Cresson mexicanus Cresson, 1878:187 P.H. WILLIAMS Bombus (Fyv.) brevivillus Franklin brevivillus Franklin, 1913:119 ?abditus (Tkalcit, 1966:271 [Megabombus})) @ TAXONOMIC STATUS. The single known female of B. abditus was described as originating from “Rep. de Guinée Beyla’ (equatorial Africa). However, it is indistinguishable from B. brevivillus according to Sakagami (1976:427) and probably represents an in- troduced or mislabelled individual (Michener, 1979). Subgenus SENEXIBOMBUS Frison Bremus (Senexibombus) Frison, 1930:3, type-species Bombus senex Vollenhoven by original designation [Bombus (Senecibombus) Kruseman, 1952:101 incorrect subsequent spelling] Bombus (Senexibombus) Richards, 1968:217 Bombus (Sx.) kulingensis Cockerell kulingensis Cockerell, 1917:266 tajushanensis Pittion1, 1949:244 Bombus (Sx.) bicoloratus Smith bicoloratus Smith, 1879:132, examined Bombus (Sx.) senex Vollenhoven Senex Vollenhoven, 1873:229 Bombus (Sx.) irisanensis Cockerell irisanensis Cockerell, 1910a:416, examined CHECKLIST OF BUMBLE BEES Subgenus DIVERSOBOMBUS Skorikov Bombus (Diversobombus ) Skorikov, 1914c:406, type-spe- cies Bombus diversus Smith by subsequent designation of Sandhouse, 1943:546 Diversibombus Skorikov, 1938b:2, unjustified emenda- tion Bombus (Dy.) trifasciatus Smith trifasciatus Smith, 1852a:43, examined montivagus Smith, 1878:168, examined montivagus Smith, 1879:131, redescribed ?wilemani Cockerell, 1911:100, examined albopleuralis Friese, 1916:108, examined ?maxwelli Pendlebury, 1923:67, examined mimeticus Richards, 1931b:529, examined malaisei (Skorikov, 1938b:2 [Diversibombus]) not of Bischoff, 1930:4 (= B. sporadicus Nylander) atropygus (Tkalcti, 1989:58 |Megabombus]}) examined @ TAXONOMIC sTaTUS. Several of these nominal taxa have been treated as separate species, for example as B. albopleuralis (= B. mimeticus) {Himalaya}, B. montivagus {northern Burma to southern China], B. maxwelli [Peninsular Malaysia] and B. wilemani [Tai- wan] (Tkalcii, 1968b, 1989). However, aside from differences in colour pattern (Fig. 13), they are closely similar in morphology and show a range of variation (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of varia- tion, I shall continue to treat them as parts of a single variable species. Bombus (Dyv.) longipes Friese longipes Friese, 1905:511 hummeli Bischoff, 1936:18, examined Bombus (Dy.) diversus Smith diversus Smith, 1869:207, examined tersatus Smith, 1869:207, examined OQ NOMENCLATURE. Tkalcti (1965) first explicitly regarded B. diversus and B. tersatus as conspecific and, following the Principle of First Reviser (ICZN, 1985: Article 24), chose B. diversus as the valid name for the species. 115 Bombus (Dv.) ussurensis Radoszkowski Ussurensis Radoszkowski, 1877b:196 {ussuriensis Morawitz, 1881:254, incorrect subsequent spelling] Subgenus MEGABOMBUS Dalla Torre Bombus (Megabombus) Dalla Torre, 1880:40, type-spe- cies Bombus ligusticus Spinola (= Bombus argillaceus (Scopoli)) by monotypy Bombus (Megalobombus) Schulz, 1906:267, unjustified emendation Bombus (Hortobombus) Vogt, 1911:56, type-species Apis hortorum Linnaeus (= Bombus hortorum (Linnaeus)) by subsequent designation of Sandhouse, 1943:559 Bombus (Odontobombus) Kriiger, 1917:61,65 (proposed as a section name but stated by Milliron, 1961:53, to be equivalent to his concept of the subgenus Megabombus Dalla Torre), type-species Apis argillacea Scopoli (= Bombus argillaceus (Scopoli)) by subsequent designa- tion of Williams, 1995:339 [Nortobombus Skorikoy, 1922b:map 3, incorrect subse- quent spelling] Hortibombus Skorikov, 1938a:146, unjustified emenda- tion Bombus (Mg.) supremus Morawitz supremus Morawitz, 1886:196 linguarius Morawitz, 1890:351 Bombus (Mg.) gerstaeckeri Morawitz Gerstdckeri Morawitz, 1881:242 Gerstaeckeri Hoffer, 1883:55, mandatory correction (ICZN, 1985: Article 32d) Bombus (Mg.) consobrinus Dahlbom consobrinus Dahlbom, 1832:49 Bombus (Mg.) tichenkoi (Skorikoy) [tichenkoi (Skorikoy, 1922a:156 [Hortobombus]) published without description] tichenkoi (Skorikov, 1925:115 [Hortobombus]) 116 ?yezoensis Matsumura, 1932:pl. 1 ?przewalskiellus (Skorikov, 1933a:59 [Hortobombus]) kurilensis Sakagami, 1954:92 @ TAXONOMIC STATUS. B. tichenkoi and B. yezoensis have apparently been regarded both as conspecific and as separate species. Sakagami (1954) described kurilensis as a (semi- melanic) subspecies of what he called ‘B. tersatus’ (a misidentification, = B. diversus). Following Tkalcit (1962), Sakagami subsequently identified his ‘B. tersatus’ as B. yezoensis (Ito & Sakagami, 1980). In this later paper he went on to recognise kurilensis as conspecific with B. tichenkoi, but then, despite having regarded kurilensis as a subspecies of what he now believed to be B. yezoensis in the earlier paper, listed B. tichenkoi as a species separate from B. yezoensis. The apparent contradiction was not explained, al- though morphological comparisons between these taxa, B. argillaceus and B. sushkini were tabulated (which show primarily that B. argillaceus 1s very different). He even noted the allopatric distributions of B. tichenkoi and B. yezoensis between the northern and southern Kurile Islands and the ‘resemblance of tichenkoi and dark individuals of yezoensis, especially in workers.’ Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat B. tichenkoi and B. yezoensis as parts of a single variable species. Bombus (Mg.) sushkini (Skorikov) [saltuarius (Skorikov, 1922a:156 [Hortobombus]) pub- lished without description] sushkini (Skorikoy, 1931:235 [Hortobombus]) examined saltuarius (Skorikov, 1931:235 |Hortobombus]) @ TAXONOMIC STATUS. B. sushkini and B. saltuarius have been regarded both as conspecific (Bischoff, 1936) and as separate species (Skorikov, 1931; Tkalct, 1974a). I have as yet seen no evidence that more than one species is involved. More evidence is awaited. O NOMENCLATURE. Skorikov (1931) provided the first valid publication of the names B. sushkini and B. saltuarius as two separate species. Subsequently, Bischoff (1936) regarded the two as conspecific and, following the Principle of First Reviser (ICZN, 1985: Article 24), chose B. sushkini as the valid name for the species. Bombus (Mg.) portchinsky Radoszkowski Portchinskij Radoszkowski, 1883:208 Portchinsky Radoszkowski, 1883:208[210], 209[211], 210[212] O NOMENCLATURE. Radoszkowski (1883) pub- lished two different spellings of B. portchinsky, P.H. WILLIAMS repeating this second form several times (page num- bers 207 and 208 are repeated twice for different pages, so the two spellings do not occur on the same page). Precedence of the correct original spelling should be determined using the Principle of the First Reviser (ICZN, 1985: Article 24), but to date I have not found an author who has correctly cited both names and then chosen one in precedence to the other (there are many incorrect subsequent spellings, e.g. Dalla Torre, 1896). Since the second form of the name has been used more recently (e.g. Baker, 1996b), I suggest that it should be given precedence. This form is a simple noun in apposition and so retains the same ending whatever the gender of the generic name with which it is combined (ICZN, 1985: Article 31b(11)). Bombus (Mg.) hortorum (Linnaeus) hortorum (Linnaeus, 1761:424 [Apis]) examined meridionalis Dalla Torre, 1879:13 hispanicus Pittioni, 1939c:244, not of Friese, 1911:571 (= B. monticola Smith) asturiensis (Tkalct, 1975:181 [Megabombus]) replace- ment name for hispanicus Pittioni, 1939c:244 ?reinigiellus (Rasmont, 1983:43 [Megabombus]) @ TAXONOMIC STATUS. The SpanishB. asturiensis has been considered a separate species from B. hortorum by Rasmont (1983, 1988), although they have been treated as conspecific by Pittioni (1939c), Tkalcti (1975), Ornosa (1986a, 1986b, 1991), Castro (1988, 1993) and, more recently, by Rasmont ef al. (1995). The two taxa are closely similar. The Spanish B. reinigiellus has also been consid- ered both as conspecific with B. hortorum (Castro, 1987) and as a separate species (e.g. Rasmont, 1983; Castro, 1988; Ornosa, 1991).The two taxa are allopatric (Rasmont, 1983), with B. reinigiellus being narrowly restricted to the Sierra Nevada of Spain, possibly as a disjunct peripheral population. B. reinigiellus is closely similar to B. hortorum, although subtle differences in characters of colour and morphology have been de- scribed (e.g. Rasmont, 1983; Castro, 1988; Ornosa, 1991). From the material I have examined, the mor- phological differences appear to be analogous to the variation between mainland and island populations of B. terrestris (see the comments on B. terrestris). Depending upon the species concept embraced, such subtle differences as those between B. reinigiellus and B. hortorum might be expected even within a single population and [| shall treat all three taxa as conspecific for the present. More evidence is awaited. COMMENT. B. hortorum has been introduced into New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr, 1995) (see the comments on B. ruderatus, B. subterraneus and B. terrestris). It occurs in Iceland, where it has also probably been introduced (Prys- Jones et al., 1981) (see the comments on B. Jucorum). CHECKLIST OF BUMBLE BEES Bombus (Mg.) argillaceus (Scopoli) Argillacea (Scopoli, 1763:305 [Apis]) ligusticus Spinola, 1806:29 ?Bombus (Mg.) ruderatus (Fabricius) ruderata (Fabricius, 1775:380 [Apis]) examined Perniger (Harris, 1776:131 [Apis]) villarricaensis Asperen de Boer, 1992a:133 @ TAXONOMICSTATUS. B. argillaceus and B. rude- ratus are similar in most characters and differ principally in the colour patterns of the queens (e.g. Reinig, 1939; Fig. 11). Scholl, Obrecht & Zimmer- mann (1992) found that hybrid queens between B. argillaceus and B. ruderatus do occur in parts of southeastern France, but are very rare. Whether or not the taxa on either side of this hybrid zone are consid- ered to be separate species therefore depends on which species concept is preferred. Because Scholl, Obrecht & Zimmermann (1992) estimated that only slight gene flow is occurring, I shall continue to treat them as separate species. COMMENT. This species has been introduced into New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr, 1995) (see the comments on B. hortorum, B. subterraneus and B. terrestris) and Chile (Arretz & Macfarlane, 1982; Asperen de Boer, 1993b). B. ruderatus also occurs on the Azores (which have never had a continental connection), where it may be pre- sumed to be an introduction (Yarrow, 1967). Bombus (Mg.) czerskii Skorikov czerskii Skorikov, 1910b:413, examined Bombus (Mg.) koreanus (Skorikov) koreanus (Skorikoy, 1933a:59 [Hortobombus]) pekingensis Bischoff, 1936:21, examined ?notocastaneus Tkalcit, 1961a:52 (provisional synonym) @ TAXONOMIC STATUS. B. notocastaneus was de- scribed from a single male from Hubei. From the description, it appears most likely to be conspecific with B. koreanus. Bombus (Mg.) melanopoda Cockerell melanopoda Cockerell, 1910a:416, examined @ TAXONOMIC STATUS. B. melanopoda is known from a single female specimen (labelled ‘Sumatra’, reverse *92.182.’) in the NHM collection in London. According to the accessions catalogue, the number on the label of this specimen refers to 8 Hymenoptera presented in November 1892 by H. O. Forbes. The area of origin is given as Borneo, although this has 117 been crossed out and Sumatra added. Forbes’ (1885) account of his travels of 1878-1883 in Indonesia shows that he did not visit Borneo, although he did visit the mountains of southern Sumatra. There is no direct account of the collection of this specimen, although he recorded bees from at least three possible localities at higher altitudes: first, in late 1880 he climbed Gunung Tenggamus, where he recorded (p. 159) ‘a few bees’ at 7200 ft (2160 m); second, in 1881 he visited Gunung Dempa, where he recorded (p. 208) ‘a fine grey-haired humble-bee (Bombus senex)’ (iden- tification by Forbes) between 7000-7700 ft (2100-2310 m); and later in the same year, near the summit of Gunung Kaba (1983 m), he recorded again (p. 228) ‘A large humble-bee (Bombus senex)’. B. melanopoda appears to be a morphologically distinct species. It can be distingished from the only other long-tongued bumblebee species known from Sumatra, B. senex, by the longer oculo-malar area of B. melanopoda, which is more than 1.5 times longer than the basal breadth of the mandible. Otherwise, the most closely related long-tongued bumble bee in any neighbouring area is B. trifasciatus from the Cameron Highlands of Peninsular Malaysia. How- ever, like most queens of the subgenus Megabombus, the holotype of B. melanopoda is easily distin- guished by its narrow longitudinal median groove subapically on gastral tergum VI (for B. trifasciatus this area is uniformly convex). The colour pattern of the B. melanopoda female is predominantly black, but the hairs of terga [V—V are very slightly paler, so there is some similarity to the darkest queens of B. koreanus, which have this pubescence brownish cream. However, unlike the few queens of B. koreanus available to me, the type of B. melanopoda has the unpunctured areas around the ocelli extend- ing to less than half the ocello-ocular distance; the dorsal furrow of the gena (between the vertex and the post-ocular area) is strongly marked anteriorly; and the dorsal face of the labral tubercles (the face adjacent to the clypeus) is more sharply separated from the anterior ventral face and more densely marked by moderate-sized punctures. Since no further individuals have been found, the possibility that the holotype of B. melanopoda is a mislabelled melanic specimen of another species of the subgenus Megabombus ought to be explored, per- haps initially through a morphometric analysis. COMMENT. Like most other Sumatran bumble bees (with the notable exception of the extensively greyish- white queens of B. senex, see Sianturi et al., 1995), the holotype of B. melanopoda is almost entirely black. It is likely that females of this species would be particu- larly easily mistaken for black individuals of B. senex (although males of B. melanopoda might be paler, as for B. koreanus). 118 Bombus (Mg.) securus (Frison) securus (Frison, 1935:346 [Bremus]) examined yuennanicus Bischoff, 1936:23, examined Bombus (Mg.) religiosus (Frison) religiosus (Frison, 1935:344 [Bremus]) examined Subgenus RHODOBOMBUS Dalla Torre Bombus (Rhodobombus) Dalla Torre, 1880:40, type-spe- cies Bremus pomorum Panzer (= Bombus pomorum (Panzer)) by subsequent designation of Sandhouse, 1943:596 Bombus (Pomobombus) Kriiger, 1917:65, type-species Bremus pomorum Panzer (= Bombus pomorum (Pan- zer)) by subsequent designation of Sandhouse, 1943:589 Pomibombus Skorikov, 1938a:145, unjustified emenda- tion Bombus (Rh.) armeniacus Radoszkowski armeniacus Radoszkowski, 1877b:202 Bombus (Rh.) mesomelas Gerstaecker® Agrorum (Schrank, 1781:397 [Apis]) arvenjfis (Gmelin in Linnaeus, 1790:2786 [Apis]) unjusti- fied replacement name for agrorum Schrank, 1781:397 mesomelas Gerstaecker, 1869:321 O NOMENCLATURE. Warncke (1986) listed B. agrorum (Schrank) as questionably conspecific with B. distinguendus, but without any explanation. Al- though I know of no extant type specimens, Schrank’s (1781) description of his B. agrorum of ‘Habitat rurv from Austria appears to me to be almost certainly of the same species as B. mesomelas, because the head is described as black and the pale hairs of the thorax and of gastral tergum I are described as grey, with the remainder of the gaster rusty or tawny-yellow (the head and the pale pubescence of the thorax and gaster are more uniformly dull yellowish fa B. distinguendus). See the comments onB. distinguendus. & APPLICATION TO ICZN. Although B. agrorum is the oldest available name for the present interpretation of this species, the name B. mesomelas has been in P.H. WILLIAMS common use for the species since 1947 (e.g. Tkalcii, 1969, 1975; Delmas, 1976; Reinig, 1974, 1981; Ozbek, 1983; Rasmont, 1983; Ornosa, 1986a, b; Rasmont et al., 1987, 1995). I know of no publications using the name B. agrorum (Schrank) since 1947, although the name B. agrorum (Fabricius) was in widespread use for another species (= B. pascuorum) until Richards (1968). It is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress the unused senior synonym, B. agrorum, and its unjustified re- placement name, B. arvensis (ICZN, 1985: Article 79) (see the comments on B. muscorum). Bombus (Rh.) pomorum (Panzer) pomorum (Panzer, 1805(86):18 [Bremus]) Lefebvrei Lepeletier, 1836:461 Subgenus KALLOBOMBUS Dalla Torre Bombus (Kallobombus) Dalla Torre, 1880:40, type-spe- cies Apis soroeensis Fabricius (= Bombus soroeensis (Fabricius)) by subsequent designation of Sandhouse, 1943:561 Bombus (Callobombus) Dalla Torre, 1896:503, unjustified emendation Bombus (Soroeensibombus) Vogt, 1911:63, type-species Apis soroeensis Fabricius (= Bombus soroeensis (Fabricius)) by monotypy [Bombus (Soroensibombus) Ball, 1914:78, incorrect sub- sequent spelling] [Sorocoénsibombus Skorikoy, 1922a: map 15, incorrect subsequent spelling] Bombus (KI.) soroeensis (Fabricius) @ Cardui (Miller, 1776:165 [Apis]) Joroeenfis (Fabricius, [1777, see Baker, 1996a:9]:246 [Apis}) Proteus Gerstaecker, 1869:325 perplexus Radoszkowski, 1884:82, not of Cresson, 1863:91 (= B. perplexus Cresson) Radoszkowskyi Dalla Torre, 1890:139, replacement name for perplexus Radoszkowski, 1884:82 miniatocaudatus Vogt, 1909:56 © NOMENCLATURE. Baker(1996a) has established CHECKLIST OF BUMBLE BEES that the name B. cardui has narrow priority over B. soroeensis by publication date. © APPLICATION TOICZN. AlthoughB. cardui is the oldest available name for the present interpretation of this species, the name B. soroeensis has been in com- mon use for the species since 1947 (e.g. Tkalct, 1969, 1975; Lgken, 1973; Alford, 1975; Delmas, 1976; Pekkarinen, 1979; Reinig, 1981; Ozbek, 1983; Rasmont, 1983; Ito, 1985; Ornosa, 1986a; Rasmont et al., 1995). The only publications using the name B. cardui since 1947 are those of Baker (1996a, b). Using this name contrary to the purpose of priority is not accepted as usage in the sense of the Code (ICZN, 1985: Article 23b), and so cannot justify the continued use of the name B. cardui in place of B. soroeensis. It is suggested that, in the interests of stability, an appli- cation be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). Subgenus ALPINOBOMBUS Skorikoy Alpinobombus Skorikov, 1914a:123, type-species Apis alpina Linnaeus (= Bombus alpinus (Linnaeus)) by subsequent designation of Frison, 1927:66 Bombus (Alpinobombus) Kriiger, 1917:62 Alpinibombus Skorikoy, 1937:53, unjustified emendation COMMENT. Species of the subgenus Alpinobombus make up the most northerly distributed of all bee faunas (e.g. K. W. Richards, 1973). Indeed, three of the five species have a nearly circumpolar distribution, as a major component of an Arctic bumble bee fauna (Williams, 1996b). This relative homogeneity of the Arctic fauna among northern continents resembles the pattern in the Arctic flora, which shows little regional differentiation in comparison with more southern flo- ras (Hooker, 1861; Walker, 1995). Bombus (Al.) hyperboreus Schonherr Arctica (Quenzel in Acerbi, 1802:253 [Apis]) hyperboreus Schénherr, 1809:57, unjustified replacement name for arcticus Quenzel, 1802:253 clydensis Yarrow, 1955:151, examined 119 @ TAXONOMIC STATUS. The identity of B. arcticus (Quenzel) has been uncertain. Warncke (1986) listed B. arcticus (Quenzel) as conspecific withB. lapponicus without any explanation. Presumably this was because B. lapponicus is extensively pale on the dorsum, al- though the pale pubescence is differentiated into yellow and red areas and much of it is much paler than Quenzel’s description. | agree with Lgken (1973) that, from the original description and the illustration (no type specimen is known to exist), B. arcticus (Quenzel) is most likely to be conspecific with B. hyperboreus, which has the pale pubescence uniformly brownish yellow. OQ NOMENCLATURE. The name B. arcticus has rarely been used for this species in preference to B. hyperboreus, and perhaps only as a misidentification of B. arcticus Kirby (see e.g. Franklin, 1913; Richards, 193 1a). Loken (1973) considered B. arcticus (Quenzel) to be a nomen oblitum, so she continued to use the name B. hyperboreus. However, nomina oblita are not supported for a publication of this date by the present Code (ICZN, 1985: Article 79c(ii1)), although it does allow that B. arcticus (Quenzel) could be suppressed by use of the Plenary Power. See the comments on B. polaris Curtis. € APPLICATION TO ICZN. Although B. arcticus is the oldest available name for the present interpretation of this species, the name B. hyperboreus has been in common use for the species since 1947 (e.g. Loken, 1973; Milliron, 1973a; K.W. Richards, 1973; Svensson & Lundberg, 1977; Hurd, 1979; Pekkarinen, 1979; Pekkarinen ef al., 1981; Reinig, 1981; Rasmont, 1983; Pekkarinen & Teras, 1993). It is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79), in order to confirm usage of B. hyperboreus as the valid name (see the comments on B. muscorum). COMMENT. B. hyperboreus has been suggested to be a social parasite in colonies of B. polaris, at least facultatively in some parts of its range (Milliron & Oliver, 1966; Loken, 1973: K. W. Richards, 1973). See the comments on Psithyrus and B. inexspectatus. Bombus (Al.) balteatus Dahlbomes balteatus Dahlbom, 1832:36 nivalis Dahlbom, 1832:40 tricolor Dahlbom, 1832:41 ?Kirbiellus Curtis in Ross, 1835:1xii kirbyellus Dalla Torre, 1896:527, unjustified emendation tristis Sparre-Schneider in Friese, 1902:495, not of Seidl, 1837:69 (= B. humilis Iliger) @ TAXONOMIC status. B. balteatus and B. kirbi- ellus have been considered conspecific by most authors (e.g. Thomson, 1872; Richards, 1931a; Skorikov, 1937; 120 Pittioni, 1942; Loken, 1973; Hurd, 1979; Thorp et al., 1983), although Milliron (1973a) considered them to be separate species that co-occur in some areas, par- ticularly in Alaska. Milliron (1973a) described several characters by which to discriminate B. balteatus and B. kirbiellus, placing particular emphasis on the shape of male gastral sternum VIII and the female malar area. From the small samples I have examined, I have been unable to find convincing evidence of discrete differences in these characters. Until more evidence to the contrary is available from critical studies of pat- terns of variation, I shall treat them as parts of a single variable species. O NOMENCLATURE. Richards (193 1a) believed B. balteatus, B. nivalis and B. tricolor to be conspecific and selected the name B. balteatus to have precedence because it was published on an earlier page (page priority is not a mandatory part of the Code, only a recommendation, see ICZN, 1985: Recommendation 24A). However, Thomson (1872:35) had already cho- sen the name B. nivalis in precedence to B. balteatus and, following the Principle of the First Reviser (ICZN, 1985: Article 24), Thomson’s action should now stand. Consequently, the valid name for this species is B. nivalis, although the Code (ICZN, 1985) allows that this name could be suppressed by use of the Plenary Power. € APPLICATIONTOICZN. AlthoughB. nivalis is the valid name for this species, the name B. balteatus has been in common use for the species since 1947 (e.g. Loken, 1973; Milliron, 1973a; Plowright & Stephen, 1973; Hurd, 1979; Pekkarinen, 1979; Reinig, 1981; Rasmont, 1983; Thorp er al., 1983; Laverty & Harder, 1988; Pekkarinen & Teras, 1993). Itis suggested that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the unused name (ICZN, 1985: Article 79) (see the comments on B. muscorum). However, the consequence of this action would be that nivalis Dahlbom would no longer be available for a subspecies of B. balteatus. Bombus (Al.) neoboreus Sladen strenuus Cresson, 1863:102, not of Harris, 1776:131 (=B. lapidarius (Linnaeus)) neoboreus Sladen, 1919:28 O NOMENCLATURE. B. strenuus Cresson (1863) is a junior secondary homonym in Bombus of Apis strenuus Harris (1776), and therefore the name B. strenuus Cresson is invalid (ICZN, 1985: Article 57c). For this species, the oldest available name is B. neoboreus, which becomes the valid name. The only publications using the name B. strenuus Cresson since 1947 of which I am aware are by Hurd (1979), Milliron (1973a) and Poole (1996), so this change of valid name is not a serious disruption of common usage. P.H. WILLIAMS Bombus (AL) polaris Curtis Arcticus Kirby in Parry, 1824:ccexvi, examined, not of Quenzel in Acerbi, 1802:253 (= B. hyperboreus Sch6nherr) Polaris Curtis in Ross, 1835:1xiii, examined diabolicus Friese, 1911:571 alpiniformis Richards, 1931a:13 © NOMENCLATURE. Loken (1973) used the name B. arcticus Kirby for this species because she consid- ered B. arcticus (Quenzel) to be a nomen oblitum. However, this is not supported by the present Code for a publication of this date (ICZN, 1985: Article 79c(iii)), although it does allow B. arcticus (Quenzel) to be suppressed by use of the Plenary Power. See the comments on B. hyperboreus. APPLICATION TO ICZN. It is suggested above that, in the interests of stability ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress B. arcticus (Quenzel), the unused senior synonym (ICZN, 1985: Article 79) of B. hyperboreus. This would free B. arcticus Kirby from junior primary homonymy with B. arcticus (Quenzel) (ICZN, 1985: Article 57b), so that it would become the valid name for this species (see the comments on B. hyperboreus). However, although the name B. arcticus Kirby has been in use for this species (e.g. Loken, 1973; Sakagami, 1976; Svensson & Lundberg, 1977; Reinig, 1981), the more frequently used name has been B. polaris (e.g. Milliron & Oliver, 1966; Milliron, 1973a: K.W. Richards, 1973; Hurd, 1979; Pekkarinen, 1979; Pekkarinen ef al., 1981; Rasmont, 1983; Pekkarinen & Teris, 1993). In the interests of stability, the application to ICZN might be extended to suppress B. arcticus Kirby, in order to conserve the current usage of B. polaris as the valid name. Bombus (AI.) alpinus (Linnaeus) alpina (Linnaeus, 1758:579 [Apis]) examined Subgenus SUBTERRANEOBOMBUS Vogt Bombus (Subterraneobombus) Vogt, 1911:62, type-spe- ciesApis subterranea Linnaeus (=Bombus subterraneus (Linnaeus)) by subsequent designation of Frison, 1927:68 CHECKLIST OF BUMBLE BEES Subterraneibombus Skorikov, 1938a:145, unjustified emendation Bombus (St.) melanurus Lepeletier melanurus Lepeletier, 1836:469, examined ?difficillimus Skorikov, 1912:609, examined subdistinctus Richards, 1928a:333, examined @ TAXONOMIC sTaTUS. Several of these nominal taxa have been treated as separate species and at least B. difficillimus may indeed prove to be a separate species. However, aside from differences in colour pattern, they are closely similar in morphology with a range of variation (Williams, 1991). Until more evi- dence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (St.) fragrans (Pallas) fragrans (Pallas, 1771:474 [Apis]) mongol Skorikoy, 1912:607, examined ?charharensis Yasumatsu, 1940:94 (provisional synonym) @ TAXONOMIC STATUS. This taxon is interpreted here in the broadest sense, to include a complex of poorly-known taxa (Williams, 1991). More evidence is awaited. Bombus (St.) amurensis Radoszkowski Amurensis Radoszkowski, 1862:590, examined @ TAXONOMIC sTaTUS. I have seen no males of this species and its precise relationships remain un- clear. Bombus (St.) fedtschenkoi Morawitz Fedtschenkoi Morawitz in Fedtschenko, 1875:5 Bombus (St.) personatus Smith personatus Smith, 1879:132, examined Roborowskyi Morawitz, 1886:197, examined Bombus (St.) subterraneus (Linnaeus) fubterranea (Linnaeus, 1758:579 [Apis]) examined Nemorum (Scopoli, 1763:307 [Apis]) ? faltuum (Panzer, 1801(75):21 [Apis]) O NOMENCLATURE. Loken (1984) interpreted B. saltuum as being conspecific with B. barbutellus (see the comments on B. barbutellus). Warncke (1986) interpreted B. saltuum as having been described from a male (presumably because the antennae were de- scribed as rather long) conspecific with B. subterraneus. No type specimen is known. The de- 121 scription of the anterior part of the gaster of B. saltuum as ashen and the middle part as nearly bald is perhaps slightly closer to B. subterraneus, because although both species may have gastral tergum I with pale hair and terga I-III sparsely haired, this hair is much shorter on B. subterraneus and the posterior fringing hairs of the terga often appear greyish-white. If B. saltuum were not accepted as most likely to be conspecific with B. subterraneus, then further action would be required. If both interpretations were con- sidered to remain supportable, then it might be considered appropriate (ICZN, 1985: Article 75b) to designate a specimen of B. subterraneus as neotype of Apis saltuum in order to conserve the current usage of B. barbutellus. Alternatively, if B. saltuum were considered more likely to be conspecific with B. barbutellus, then B. saltuum would become the oldest available name for that species, even though the name has not been used in the last 50 years. In the interests of stability (ICZN, 1985: Article 23b), an application could then be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79). COMMENT. This species has been introduced into New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr, 1995). Bombus (St.) distinguendus Morawitz nemorum (Fabricius, 1775:382 [Apis]) not of Scopoli, 1763:307 (= B. subterraneus (Linnaeus)), not of Fabricius, 1775:380 (?= B. bohemicus Seidl) elegans Seidl, 1837:67 distinguendus Morawitz, 1869:32 O NOMENCLATURE. The nameB. elegans has been applied to several taxa by different authors. Tkalcit (1969:901-903) reasoned that Seidl had originally described B. elegans from an individual of the species that has more recently been known by the name B. distinguendus, although Seidl’s original type is lost. According toTkalcti, a specimen of B. mesomelas may then have been substituted as the type, but now this cannot be found either. Any remaining confusion could be resolved by the designation of an appropriate neotype. See the comments on B. mesomelas. € APPLICATIONTOICZN. AlthoughB. elegans may be the oldest available name for the present interpreta- tion of this species, the name B. distinguendus has been in common use for the species since 1947 (e.g. Tkalcti, 1969, 1974a; Loken, 1973; Alford, 1975; Delmas, 1976; Sakagami, 1976; Pekkarinen, 1979; Reinig, 1981; Pekkarinen er al/., 1981; Rasmont, 1983; Pekkarinen & Teréas, 1993; Rasmont ef al., 1995). I know of no publications using the name B. elegans for this taxon (only for B. mesomelas Gerstaecker as a misidentification) since 1947. It is suggested that, in 122 the interests of stability (ICZN, 1985: Article 23b), and to prevent confusion with B. mesomelas, an appli- cation be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). Bombus (St.) appositus Cresson appositus Cresson, 1878:183 Bombus (St.) borealis Kirby borealis Kirby, 1837:272 Subgenus ALPIGENOBOMBUS Skorikov Alpigenobombus Skorikov, 1914a:128, type-species Alpigenobombus pulcherrimus Skorikov (= Bombus kashmirensis Friese) by subsequent designation of Williams, 1991:65 Bombus (Mastrucatobombus) Kriger, 1917:66, type-spe- cies Bombus mastrucatus Gerstaecker (= Bombus wurflenii Radoszkowski) by monotypy Bombus (Alpigenobombus) Frison, 1927:64 [Nobilibombus Skorikov, 1933a:62, published without fixa- tion of type-species] [Bombus (Nobilibombus) Bischoff, 1936:12, type-species Nobilibombus morawitziides Skorikov (= Bombus nobilis Friese) by monotypy, published as a junior synonym] Alpigenibombus Skorikov, 1938b:1, unjustified emenda- tion [Pyrobombus (Nobilibombus) Milliron, 1961:54, type-spe- cies Bombus nobilis Friese (cited as Bombus nobilis Skorikov) by original designation, published as a junior synonym] Bombus (Nobilibombus) Richards, 1968:222, type-species Bombus nobilis Friese by original designation (see Williams, 1991) [Alpegenobombus Wang, 1979:188, incorrect subsequent spelling] @ TAXONOMIC sTATUS. Richards (1968) treated Alpigenobombus and Nobilibombus as separate subgenera. Following Bischoff (1936), I have treated them as a single subgenus Alpigenobombus (Williams, 1991), for which the evidence for monophyly is strong (Williams, 1995). P.H. WILLIAMS Bombus (Ag.) kashmirensis Friese kashmirensis Friese, 1909[September, Tkalcii, 1974b]:673 examined stramineus Friese, 1909[September, Tkalct, 1974b]:673 tetrachromus Cockerell, 1909[November, Tkalci, 1974b]:397, examined pulcherrimus (Skorikov, 1914a:128 [Alpigenobombus]) @ TAXONOMIC sTaTUS. Several of these nominal taxa have been treated as separate species. However, aside from differences in colour pattern, they are closely similar in morphology with a range of varia- tion (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. O NOMENCLATURE. Tkalcti (1974b) first regarded B. kashmirensis and B. stramineus as conspecific and, following the Principle of First Reviser (ICZN, 1985: Article 24), chose B. kashmirensis as the name for the species. Bombus (Ag.) wurflenii Radoszkowski Wurflenii Radoszkowski, 1859:482, examined [Wurfleini Radoszkowski, 1877b:191, incorrect subsequent spelling] mastrucatus Gerstaecker, 1869:326, examined alpigenus Morawitz, 1874:132 Bombus (Ag.) nobilis Friese ?validus Friese, 1905:510, examined (provisional syno- nym) nobilis Friese, 1905:513 ?sikkimi Friese, 1918:82, examined (provisional syno- nym) [morawitziides Skorikov, 1922a:159, published without description] [moravitziides Skorikov, 1931:203, published without de- scription] ?morawitziides (Skorikov, 1933a:62 [Nobilibombus]) ex- amined (provisional synonym) Dxizangensis Wang, 1979:188, examined (provisional syno- nym) chayaensis Wang, 1979:189, examined, new synonym @ TAXONOMIC STATUS. Several of these nominal taxa have been treated as separate species. Friese (1905) described B. validus initially (p. 510) as having a quadrate malar area and untoothed mandi- bles, but went on (p. 517) to place it within the mastrucatus (= B. wurflenit) group, which he charac- terised as having a short malar area and toothed mandibles. Tkalcii (1987) designated as lectotype of B. validus a female with a quadrate malar area and multi-toothed mandibles. He also synonymised B. morawitziides with B. validus. The type specimens of B. nobilis have also been in some doubt (Richards, 1968). In the same publication CHECKLIST OF BUMBLE BEES as the description of B. validus, Friese (1905) de- scribed the female of B. nobilis as having a‘quadratisch’ malar area and 4—5 teeth on the mandible (even though he placed it [p. 519] ina group withB. lapidarius).The original description lists several females (particularly from Sichuan), but the only putative type female that I have been able to examine (although it carries no Friese ‘type’ label) is in the Berlin museum collection and is a specimen of B. friseanus labelled ‘Kashgar’ (this locality is outside the known distribution range of either B. nobilis or B. friseanus). The specimen does not match the original description of the mandibles of B. nobilis and so cannot be considered a valid syntype. Nonetheless, the identity of B. nobilis is clear from the original description, so the designation of a neotype is not justified (ICZN, 1985: Article 75b). B. chayaensis appears to me to be very closely similar to the yellow banded B. nobilis (in the strict sense) and I am unaware of any reason to treat them as separate species. B. nobilis is interpreted here in the broadest sense, to include a complex of morphologically closely simi- lar taxa (Williams, 1991). At least some of the taxa included may prove to be separate species from B. nobilis. The most obvious variation is in the colour of the pale thoracic bands, which may be yellow (B. nobilis), yellow-white (B. sikkimi), gey-white (B. morawitziides), or almost completely replaced by black (B. validus). However, aside from these differences in colour pattern, they are similar in morphology with a range of variation. Until more evidence to the contrary is available from critical studies of patterns of varia- tion, I shall treat them as parts of a single variable species. O NOMENCLATURE. Following the Principle of First Reviser (ICZN, 1985: Article 24), and as the first author to regard these taxa as conspecific, I select the name B. nobilis as the valid name in preference to B. validus from the available names for this species from Friese (1905). Bombus (Ag.) genalis Friese genalis Friese, 1918:84, examined @ TAXONOMIC sTaTUS. Ihave seen no males of this species and its precise relationships remain unclear. Bombus (Ag.) grahami (Frison) grahami (Frison, 1933:334 [Bremus]) Bombus (Ag.) breviceps Smith nasutus Smith, 1852a:44, examined breviceps Smith, 1852a:44, examined dentatus Handlirsch, 1888:227 simulus Gribodo, 1892:114, examined 123 orichalceus Friese, 1916:107 rufocognitus Cockerell, 1922:4, examined pretiosus Bischoff, 1936:11, examined, not of Friese, 1911:571 (= B. polaris Curtis) 2angustus Chiu, 1948:59 (provisional synonym) bischoffiellus (Tkalct, 1977:224 [Alpigenobombus]) re- placement name for pretiosus Bischoff, 1936:11 @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species. At least B. dentatus [Himalaya] and B. angustus [Taiwan] may prove to be separate species (e.g. Tkalct, 1968), 1989). However, aside from differences in colour pattern, they are similar in morphology with a range of varia- tion (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. O NOMENCLATURE. Tkalcii (1968d) first regarded B. nasutus and B. breviceps as likely to be conspecific and, following the Principle of First Reviser (ICZN, 1985: Article 24), chose B. breviceps as the name for the species. Subgenus PYROBOMBUS Dalla Torre Bombus (Pyrobombus) Dalla Torre, 1880:40, type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by monotypy Bombus (Poecilobombus) Dalla Torre, 1882:23, type-spe- cies Bombus sitkensis Nylander by subsequent designation of Sandhouse, 1943:589 [Bombus (Pyrrhobombus) Dalla Torre, 1882:28, incorrect subsequent spelling] Bombus (Pyrrhobombus) Dalla Torre, 1896:503, unjusti- fied emendation Bombus (Pratobombus) Vogt, 1911:49, type-species Apis pratorum Linnaeus (= Bombus pratorum (Linnaeus)) by subsequent designation of Frison, 1927:67 [Bombus (Pratibombus) Ball, 1914:78, incorrect subse- quent spelling] Bombus (Anodontobombus ) Kriiger, 1917:61,65 (proposed as a section name but stated by Milliron, 1961:53, to be synonymous with his concept of the subgenus Pyrobombus Dalla Torre), type-species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by subse- quent designation of Williams, 1991:69 124 Bombus (Uncobombus) Vogt in Kriiger, 1917:65 (pro- posed as a group name but stated by Milliron, 1961:53, to correspond to his concept of Pyrobombus Dalla Torre), type-speciesApis hypnorum Linnaeus (=Bombus hypnorum (Linnaeus)) by subsequent designation of Williams, 1991:69 Bombus (Lapponicobombus) Quilis-Pérez, 1927:19, type- speciesApis lapponica Fabricius (=Bombus lapponicus (Fabricius)) by subsequent designation of Milliron, 1961:58 [Bombus (Hypnorubombus) Quilis-Pérez, 1927:19, incor- rect original spelling] [Bombus (Laponicobombus) Quilis-Pérez, 1927:63, incor- rect original spelling] Bombus (Hypnorobombus) Quilis-Pérez, 1927:97, type- species Apis hypnorum Linnaeus (= Bombus hypnorum (Linnaeus)) by monotypy Pratibombus Skorikoy, 1938b:1, unjustified emendation COMMENT. This is the largest subgenus of Bombus. The highest richness of Pyrobombus species occurs in the New World (there are no species known from south of Panama), although the earliest-diverging species within the subgenus appear to be in the Old World (Williams, 1991). This is the opposite pattern to that shown by the next-largest subgenus (of social para- sites), Psithyrus (see the comments on the subgenus Psithyrus). Bombus (Pr.) abnormis (Tkalci) abnormis (Tkalcii, 1968a:33 [Pyrobombus]) examined Bombus (Pr.) hypnorum (Linnaeus) Hypnorum (Linnaeus, 1758:579 [Apis]) examined leucopygus Wlliger, 1806:172 calidus Erichson in Middendorff, 1851:65 fletcheri Richards, 1934:90, examined insularis Sakagami & Ishikawa, 1969:180, not of Smith, 1861:155 (= B. insularis (Smith)) Koropokkrus Sakagami & Ishikawa, 1972:610, replace- ment name for insularis Sakagami & Ishikawa, 1969:180 @ TAXONOMIC sTaTuS. B. hypnorum is a broadly distributed species with a fairly easily recognised brown-black-white colour pattern (e.g. Reinig, 1939; Williams, 1991). Recently, Starr (1992) has described what appears to be a divergent, brownish-black or- ange-tailed colour form from a disjunct peripheral population on the island of Taiwan. Bombus (Pr.) perplexus Cresson perplexus Cresson, 1863:91 Bombus (Pr.) haematurus Kriechbaumer haematurus Kriechbaumer, 1870:157 Bombus (Pr.) subtypicus (Skorikov) leucopygus Morawitz in Fedtschenko, 1875:3, not of Illiger, P.H. WILLIAMS 1806:172 (= B. hypnorum (Linnaeus)) [leucopygos (Skorikov, 1914b:294 [Pratobombus]) incor- rect subsequent spelling] subtypicus (Skorikoy, 1914b:294 [Pratobombus]) exam- ined leucurus Bischoff & Hedicke, 1931:391, replacement name for leucopygus Morawitz in Fedtschenko, 1875:3 kohistanensis (Tkalcti, 1989:49 [Pyrobombus]) examined Bombus (Pr.) mirus (Tkalcii) mirus (Tkalcu, 1968a:37 [Pyrobombus]) examined ?tibetanus Friese, 1913:86, examined, not of Morawitz, 1886:202 (= B. tibetanus (Morawitz)) Bombus (Pr.) lemniscatus Skorikov lemniscatus Skorikov, 1912:607, examined flavopilosus Friese, 1918:84, examined peralpinus Richards, 1930:646, examined Bombus (Pr.) lepidus Skorikoy lepidus Skorikoy, 1912:606, examined genitalis Friese, 1913:85, examined tetrachromus Friese, 1918:85, examined, not of Cockerell, 1909:397 (= B. kashmirensis Friese) ?yuennanicola Bischoff, 1936:7, examined Bombus (Pr.) infirmus (Tkalcit) leucurus Bischoff, 1936:8, examined, not of Bischoff & Hedicke, 1931:391 (= B. subtypicus (Skorikov)) infirmus (Tkalct, 1968a:24 [Pyrobombus]) replacement name for /eucurus Bischoff, 1936:8 Bombus (Pr.) parthenius Richards parthenius Richards, 1934[14April, Williams & Cameron, 1993]:89, examined ?sonani (Frison, 1934[30 April, Williams & Cameron, 1993]:175 [Bremus]) examined ?infrequens (Tkalctt, 1989:56 [Pyrobombus]) examined (provisional synonym) @ TAXONOMIC STATUS. Several of these nominal taxa have been treated as separate species. At least B. sonani [Taiwan] and B. infrequens [northern Burma to southern China] may prove to be separate species. However, aside from differences in colour pattern, they are closely similar in morphology (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Pr.) luteipes Richards luteipes Richards, 1934:89, examined ?avanus (Skorikoy, 1938b:2 [Pratibombus]) (provisional synonym) signifer (Tkalctt, 1989:52 [Pyrobombus]), examined @ TAXONOMIC STATUS. The identity of B. avanus CHECKLIST OF BUMBLE BEES is in doubt because the type cannot be found. The description of the colour pattern resembles B. parthenius and B. luteipes, and the description of the longer than usual squama (= gonostylus) and the half- crooked apex of the sagitta (= penis valve) appear to be closely similar to B. luteipes, so these taxa are very likely to be conspecific. More evidence is awaited. Bombus (Pr.) flavescens Smith flavescens Smith, 1852a:45, examined mearnsi Ashmead, 1905:959 baguionensis Cockerell, 1920:631, new synonym tahanensis Pendlebury, 1923:65, examined ?rufoflavus Pendlebury, 1923:66, examined (provisional synonym) @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species. B. rufoflavus [Peninsular Malaysia] and B. baguionensis [Philip- pines] are particularly distinct in colour pattern. They may prove to be separate species, but from the material available from a few sites, they appear to me to be closely similarin morphology to B. flavescens (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Pr.) rotundiceps Friese rotundiceps Friese, 1916:108, examined montivolanoides Sakagami & Yoshikawa, 1961:431 Shillongensis (Tkalct, 1974b:334 [Pyrobombus]) exam- ined Bombus (Pr.) beaticola (Tkalcii) beaticola (Tkalcti, 1968a:28 [Pyrobombus]) examined Bombus (Pr.) picipes Richards flavus Friese, 1905:517, examined, not of Pérez, 1884:265 (= B. campestris (Panzer)) picipes Richards, 1934:90, examined klapperichi Pittioni, 1949:266, examined ?nikiforuki Tkalct, 1961b:354 (provisional synonym) @ TAXONOMIC STATUS. B. nikiforuki was described from a single worker from Qinghai. From the descrip- tion, it appears to be closely similar to B. picipes and is likely to be conspecific. O NOMENCLATURE. With Psithyrus regarded as being a subgenus of the genus Bombus (Williams, 1991, 1995), B. pratorum subsp. flavus Friese (1905) becomes a junior secondary homonym in Bombus of Psithyrus campestris var. flavus Pérez (1884) (deemed to be subspecific, see ICZN, 1985: Article 45g(ii)), and therefore the name B. flavus Friese is invalid (ICZN, 1985: Article 57c). For this species, the oldest available name of which I am aware is B. parthenius 125 var. picipes Richards, 1934 (deemed to be subspecific, see ICZN, 1985: Article 45g(11)), which becomes the valid name, B. picipes. The only publications using the name B. flavus Friese since 1947 of which I am aware are by Sakagami (1972), Ito (1993) and Yao & Luo (1997), so this change of valid name is not a serious disruption of common usage. Bombus (Pr.) ardens Smith ardens Smith, 1879:133, examined andreae Friese, 1910:405, examined Bombus (Pr.) modestus Eversmann modestus Eversmann, 1852:134 Baikalensis Radoszkowski, 1877b:203 nymphae Skorikoy, 1910b:409 eversmanni Skorikoy, 1910c:581, not infrasubspecific af- ter Skorikov, 1922a:149 Bombus (Pr.) cingulatus Wahlberg cingulatus Wahlberg, 1854:208 COMMENT. The distribution of B. cingulatus accord- ing to Reinig (1939) is shown in Fig. 10. ?Bombus (Pr.) oceanicus Friese oceanicus Friese, 1909:675, examined oceanicus Friese & Wagner, 1910:52, redescribed @ TAXONOMICSTATUS. B. oceanicus is known only from the Kurile Islands. A particularly close relation- ship with the otherwise broadly distributed B. cingulatus (absent from the Kuriles, but present in Kamchatka, Reinig, 1939; Ito & Sakagami, 1980; Fig. 10) has been suggested by Ito & Sakagami (1980) and it is possible that they are conspecific. More evidence is awaited. Bombus (Pr.) brodmannicus Vogt Brodmannicus Vogt, 1909:49, examined Bombus (Pr.) pratorum (Linnaeus) pratorum (Linnaeus, 1761:424 [Apis]) examined COMMENT. This species was deliberately introduced into Sydney, Australia, although it is not known to have persisted (Oliff, 1895). Until the twentieth cen- tury, B. pratorum was not known from Ireland, where it is now well established (see references in Alford, 1975, 1980) (see comments on B. monticola). Bombus (Pr.) jonellus (Kirby) Jonella (Kirby, 1802:338 [Apis]) examined alboanalis Franklin, 1913:385 126 @ TAXONOMIC status. B. alboanalis has been regarded both as a separate species (Franklin, 1913; Frison, 1927) and as conspecific with either B. frigidus (Burks, 1951; Hurd, 1979; Poole, 1996) or B. jonellus (Williams, 1991 [as B. jonellus from western Canada]; Scholl et al., 1995). Recently, Scholl et al. (1995) concluded from stud- ies of enzyme mobility morphs that whereas B. alboanalis and B. frigidus have separate gene pools, in contrast, B. alboanalis and B. jonellus show a low level of genetic differentiation. They also noted the lack of colour gradation between sympatric B. alboanalis and B. frigidus. From the limited amount of material I have exam- ined, I believe that B. alboanalis and B. jonellus are morphologically closely similar. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Pr.) pyrenaeus Pérez pyrenaeus Pérez, [1880, see Baker, 1996d:300]:127, not of Lepeletier, 1832:375 (= B. rupestris (Fabricius)) tenuifasciatus Vogt, 1909:49 [pyreneus Pagliano, 1995:23, incorrect subsequent spell- ing] O NOMENCLATURE. With Psithyrus regarded as being a subgenus of the genus Bombus (Williams, 1991, 1995), B. pyrenaeus Pérez (1880) becomes a junior secondary homonym in Bombus of Psithyrus pyrenaeus Lepeletier (1832), and therefore the name B. pyrenaeus Pérez is invalid (ICZN, 1985: Article 57c). The next available name, tenuifasciatus, was used by Vogt (1909) for individuals with particular colour patterns from both B. pyrenaeus Pérez and B. sichelii. The choice of which of these two homonyms should have precedence depends on the Principle of the First Reviser (ICZN, 1985: Article 24). As far as I have been able to discover, Tkalciti (1973:266) is the first author to have recognised this problem. He recog- nised precedence for B. pyrenaeus ssp. tenuifasciatus Vogt. Consequently, the oldest available name for this species, and therefore the valid name, is B. tenuifasciatus. & APPLICATION TO ICZN. Although B. tenuifasciatus is the oldest available name for this species, the name B. pyrenaeus has been in common use for the species since 1947 (e.g. Krusemen, 1958; Tkalct, 1969, 1973, 1975; Reinig, 1972, 1981; Delmas, 1976; Rasmont, 1983; Ornosa, 1986; Williams, 1991; Rasmont ef al., 1995). It is suggested that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the senior homo- nym (ICZN, 1985: Article 79) (see the comments on B. muscorum). However, the consequence of this action P.H. WILLIAMS would be that pyrenaeus (Lepeletier) would no longer be available for a subspecies of B. rupestris. Bombus (Pr.) biroi Vogt biroi Vogt, 1911:51, examined nursei Friese, 1918:84, examined Yagnatus Skorikov, 1933b:248, examined, not of Skorikov, 1912:97 (= B. monticola Smith) ?kotzschi Reinig, 1940:227, examined @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species. However, aside from differences in colour pattern, they are closely similar in morphology with a range of varia- tion (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Pr.) frigidus Smith frigidus Smith, 1854:399, examined Bombus (Pr.) sandersoni Franklin sandersoni Franklin, 1913:353 Bombus (Pr.) flavifrons Cresson pleuralis Nylander, 1848:231, examined flavifrons Cresson, 1863:105, new synonym @ TAXONOMIC STATUS. B. pleuralis and B. flavifrons were regarded as separate species by Franklin (1913), and Poole (1996) also lists them as separate species, without explanation. In my opinion, the lectotype of B. pleuralis designated by Milliron (1960:95) is an individual of the dark form of B. flavifrons (see descriptions of variation by e.g. Stephen, 1957; Thorp et al., 1983). See also the comments on B. mixtus. O NOMENCLATURE. B. pleuralisis the oldest avail- able name for this species. € APPLICATION TOICZN. Although B. pleuralis is the oldest available name for the present interpretation of this species, the name B. flavifrons has been in common use for the species since 1947 (e.g. Stephen, 1957; Thorp, 1969, 1970; Plowright & Stephen, 1973; Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright & Owen, 1980; Thorp et al., 1983; Laverty & Harder, 1988). I know of no publications using the name B. pleuralis since 1947, apart from the list by Poole (1996). It is suggested that, in the interests of stability (ICZN, 1985: Article 23b), an application be made to ICZN to use its Plenary Power to suppress the unused senior synonym (ICZN, 1985: Article 79) (see the rn CHECKLIST OF BUMBLE BEES comments on B. muscorum). However, the conse- quence of this action would be that pleuralis would no longer be available for a species or for a subspecies of B. flavifrons. Bombus (Pr.) centralis Cresson centralis Cresson, 1864:41 Bombus (Pr.) vandykei (Frison) vandykei (Frison, 1927:375 [Bremus]) cascadensis (Milliron, 1970a:382 [Pyrobombus]) Bombus (Pr.) caliginosus (Frison) caliginosus (Frison, 1927:376 [Bremus]) Bombus (Pr.) vagans Smith vagans Smith, 1854:399, examined Bombus (Pr.) mixtus Cresson® Praticola Kirby, 1837:274 mixtus Cresson, 1878:186, not of Kriechbaumer, 1870:160 (= B. maxillosus Klug), new synonym @ TAXONOMICSTATUS. The identity of B. praticola has remained uncertain (e.g. Cresson, 1863; Franklin, 1913). Recently, Poole (1996) has listed B. praticola, B. mixtus and B. flavifrons as separate species without explanation. Although I know of no type material, Kirby pro- vided a description of B. praticola from northern Canada (latitude 65° North) with a colour pattern (including anterior half of abdomen yellow, posterior ferruginous) that for individuals from this area is most likely to be conspecific either with B. mixtus (some individuals have few black hairs on gastral terga II- III), or with B. flavifrons (which has terga V—VI black, although this is not always apparent from the dorsal view). In his original description of B. flavifrons, Cresson (1863) conceded that this might be the same species as Kirby’s B. praticola, and he went on to write (p. 106) that he had not yet identified B. praticola. Franklin (1913:371) wrote that he had ‘been unable to decide whether the original description of B. praticolus [sic] referred to this species [B. flavifrons] or to the colour variant of pleuralis [intermediate colour pat- terns between B. flavifrons and B. pleuralis].’ Milliron (1971:42) subsequently listed Pyrobombus praticola flavifrons (Cr.) as a member of his ‘Praticola Group’. However, here I follow R. Miller (in Jitt.), who believes that the original material was more likely to have been of the species that has come to be known as B. mixtus. See the comments on B. flavifrons. O NOMENCLATURE. B. praticola is probably the oldest available name for this species. Any remaining 127 confusion could be resolved by the designation of an appropriate neotype (e.g. see the comments on B. subterraneus). © APPLICATION TOICZN. Although B. praticola is probably the oldest available name for this species, the name B. mixtus has been in common use for the species since 1947 (e.g. Stephen, 1957; Thorp, 1970; Plowright & Stephen, 1973; K. W. Richards, 1973; Macior, 1975; Sakagami, 1976; Hurd, 1979; Plowright & Owen, 1980; Thorp et al., 1983; Laverty & Harder, 1988; Macfarlane et al., 1994). It is suggested that, in the interests of stability, an application be made to ICZN to use its Plenary Power to suppress the senior synonym and homonym (ICZN, 1985: Article 79) (see the comments on B. muscorum). However, the conse- quence of this action would be that mixtus (Kriech- baumer) would no longer be available for a subspecies of B. maxillosus. Bombus (Pr.) sitkensis Nylander Sitkensis Nylander, 1848:235 Bombus (Pr.) melanopygus Nylander melanopyge Nylander, 1848:236 Edwardsii Cresson, 1878:184 melampygus Handlirsch, 1888:231, unjustified emenda- tion [melanopygus Viereck, 1904:99, incorrect subsequent spell- ing] melanopygus Franklin, 1913:334, justified emendation @ TAXONOMIC STATUS. B. melanopygus and B. edwardsii were shown by Owen & Plowright (1980) to differ principally by a single allele controlling the colour of the pubescence on gastral terga II-III. There can be little doubt that they are conspecific. Bombus (Pr.) lapponicus (Fabricius) lapponica (Fabricius, 1793:318 [Apis]) ?sylvicola Kirby, 1837:272 zhaosu Wang, 1985:162, examined, new synonym @ TAXONOMIC sTATUS. B. sylvicola is morpho- logically closely similar to B. lapponicus, and it has been suggested repeatedly that they may be conspecific (e.g. Sladen, 1919; Skorikoy, 1922a, 1937; Pittioni, 1942, 1943; Thorp, 1962; Thorp et al., 1983). B. zhaosu was described from material from Xinjiang, China, and is closely similar toB. lapponicus. These three nominal taxa have been treated as sepa- rate species. However, aside from differences in colour pattern, they are closely similar in morphology. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. 128 ?Bombus (Pr.) monticola Smith montanus Smith, 1844:549, not of Lepeletier, 1836:463 (= B. ruderarius (Miiller)) monticola Smith, 1849:lx, replacement name formontanus Smith, 1844:549 lugubris Sparre-Schneider, 1909:155, not of Kriechbaumer, 1870:159 (= B. maxillosus (Klug)) scandinavicus Friese, 1912:684, replacement name for lugubris Sparre-Schneider, 1909:255 @ TAXONOMIC STATUS. B. scandinavicus (= B. monticola) and B. lapponicus are names that were applied initially to two colour forms in Scandinavia. Lgken (1973) reported that these two taxa overlap narrowly in distribution and intergrade. However, they have been found to differ consistently (for samples analysed) in the composition of cephalic secretions (Bergstr6m & Svensson, 1973; Svensson & Bergstré6m, 1977). Svensson (1973, 1979) also described subtle differences in morphological characters, although other morphological studies by Lgken (1973) and Pekkarinen (1979) found no distinct differences. Pekkarinen (1982, in litt.) now believes that they are separate species. It remains possible that there is a hybrid zone where the colour forms intergrade, with some gene flow. In this case, depending on the species concept embraced, these taxa might be considered conspecific (see the com- ments on B. ruderatus). Until further evidence is avail- able, I shall continue to treat them as separate species. COMMENT. Until the twentieth century B. monticola was not known from Ireland, where it is now estab- lished (see references in Alford, 1975, 1980) (see comments on B. pratorum). Bombus (Pr.) bimaculatus Cresson bimaculatus Cresson, 1863:92 Bombus (Pr.) impatiens Cresson impatiens Cresson, 1863:90 Bombus (Pr.) vosnesenskii Radoszkowski Vosnesenskii Radoszkowski, 1862:589 Bombus (Pr.) bifarius Cresson bifarius Cresson, 1878:185 andamanus Gribodo, 1882:268, examined fernaldi Franklin, 1911:157, not a replacement name @ TAXONOMIC sTaTUS. B. andamanus was de- scribed as originating from ‘Andaman’ (= Andaman Islands, Indian Ocean), but appears to be a mislabelled queen of B. bifarius from western North America (Tkalcti, 1966). I have examined this specimen and agree with this identification (i.e. contrary to Richards, 1929, itis nota species of the subgenus Bombus s. str.). P.H. WILLIAMS Bombus (Pr.) huntii Greene Huntii Greene, 1860:172 Bombus (Pr.) ternarius Say ternarius Say, 1837:414 ornatus Smith, 1854:398, examined Bombus (Pr.) ephippiatus Say ephippiatus Say, 1837:414 formosus Smith, 1854:403, examined lateralis Smith, 1879:134, examined ?wilmatte Cockerell, 1912:21, examined ?alboniger Franklin, 1915:409, examined folsomi (Frison, 1923:322 [Bremus]) examined @ TAXONOMIC STATUS. B. wilmattae, B. alboniger and B. ephippiatus have been regarded both as conspecific and as separate species. Recently, B. wilmattae and B. ephippiatus were regarded as separate species by Labougle ef al. (1985) and Labougle (1990), who described diagnostic charac- ters of colour pattern and morphology. However, D. Yanega (in litt.) and G. Chavarria (pers. com.) believe that all of these nominal taxa are part of the wide- spread and variable B. ephippiatus. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. B. folsomi was described as originating from “Kina Bala / N. Borneo’ (= Gunung Kinabalu, Sabah), but appears to be a mislabelled queen of B. ephippiatus, probably from Costa Rica or Panama (Starr, 1989). I have examined this specimen and agree with this identification. Subgenus FESTIVOBOMBUS Tkalcii [Atrocinctob.{ombus | Skorikov, 1933b:244, published with- out description] Pyrobombus (Festivobombus) Tkalcu, 1972:26, type-spe- cies Bombus festivus Smith by original designation Bombus (Festivobombus) Williams, 1985b:240 @ TAXONOMIC STATUS. Richards (1968) treated B. atrocinctus (= B. festivus) as a species of the CHECKLIST OF BUMBLE BEES subgenus Pyrobombus, even though this required nu- merous exceptions in the diagnosis. I have followed Tkalcii(1972, 1974) in treating Festivobombus and Pyrobombus as separate subgenera (Williams, 1991), because together they do not form a monophyletic group (Williams, 1995). Bombus (Fs.) festivus Smith festivus Smith, 1861:152, examined atrocinctus Smith in Horne, 1870:193, examined terminalis Smith in Horne, 1870:193, examined Subgenus RUFIPEDIBOMBUS Skorikov Rufipedibombus Skorikov, 1922a:156, type-species Bombus rufipes Lepeletier by monotypy Bombus (Rufipedibombus) Richards, 1930:638 Bombus (Rufipedobombus) Kruseman, 1952:102, unjusti- fied emendation Bombus (Rf) rufipes Lepeletier rufipes Lepeletier, 1836:473 richardsi (Frison, 1930:6 |Bremus]) Bombus (Rf.) eximius Smith eximius Smith, 1852b:47, examined latissimus Friese, 1910:405 Subgenus PRESSIBOMBUS Frison Bremus (Pressibombus) Frison, 1935:342, type-species 129 Bremus pressus Frison (= Bombus pressus (Frison)) by original designation Bombus (Pressobombus) Kruseman, 1952:102, unjusti- fied emendation Bombus (Pressibombus) Richards, 1968:217 Bombus (Pe.) pressus (Frison) pressus (Frison, 1935:342 [Bremus]) Subgenus BOMBUS in the strict sense Bombus (Leucobombus) Dalla Torre, 1880:40, type-spe- cies Apis terrestris Linnaeus (= Bombus terrestris (Linnaeus)) by subsequent designation of Sandhouse, 1943:564 Bombus (Terrestribombus) Vogt, 1911:55, type-species Apis terrestris Linnaeus (= Bombus terrestris (Linnaeus)) by subsequent designation of Frison, 1927:67 Bombus (Bo.) sporadicus Nylander sporadicus Nylander, 1848:233 malaisei Bischoff, 1930:4 Bombus (Bo.) tunicatus Smith tunicatus Smith, 1852a:43, examined vallestris Smith, 1878:8 gilgitensis Cockerell, 1905:223, examined Bombus (Bo.) franklini (Frison) franklini (Frison, 1921:147 [Bremus]) @ TAXONOMIC STATUS. This species has been treated as conspecific with B. occidentalis (= B. terricola) by Milliron (1971), but has since been shown to be very distinct in morphology by Plowright & Stephen (1980) and Williams (1991), and in enzyme mobilities by Scholl, Thorp & Obrecht (1992). COMMENT. 8B. franklini has one of the narrowest distributions of any bumble bee species world-wide. All recent specimens have been collected within a 60 mile (38 km) radius of Grants Pass, Oregon (Thorp, 1970; Thorp et al., 1983). 130 Bombus (Bo.) affinis Cresson affinis Cresson, 1863:103 Bombus (Bo.) ignitus Smith ignitus Smith, 1869:207, examined terminalis Smith, 1873:206, examined, not of Smith in Horne, 1870:193 (= B. festivus Smith) japonicus Dalla Torre, 1890:139, replacement name for terminalis Smith, 1873:206 Bombus (Bo.) terrestris (Linnaeus) terreftris (Linnaeus, 1758:578 [Apis]) Audax (Harris, 1776:130 [Apis]) not of Harris, 1776:137 (= Anthophora sp.) canariensis Pérez, 1895:191 maderensis Erlandsson, 1979:191 @ TAXONOMICSTATUS. B. terrestris, B. maderensis and B. canariensis have been regarded both as conspecific and as separate species. Erlandsson (1979) argued that the dark individuals from the Canary Islands, previously placed within B. terrestris by for example Kriiger (1954, 1956), are a separate species, B. canariensis. Erlandsson also ar- gued that individuals from the island of Madeira, previously placed within B. terrestris by Bischoff (1937), are a separate species, B. maderensis. In both cases the morphological characters used to support these distinctions are not strongly divergent from the broad variation within B. terrestris in the broad sense. Rasmont (1984) regarded these three taxa as separate species, but Pekkarinen & Kaarnama (1994) treated them as conspecific. Recent work by Estoup et al. (1996) has found that although European mainland populations do not vary significantly among themselves in mitochondrial genes, all island populations studied (from six Medi- terranean islands in addition to B. canariensis) show significant differences from the mainland populations. Consequently, viewing these three nominal taxa as separate species may be one interpretation, but this appears to depend on adopting a species concept that admits little colour, morphological or genetic varia- tion within a species and regards current geographical isolation as highly indicative. I prefer to regard these taxa as conspecific until further evidence is available. O NOMENCLATURE. Day (1979) described how none of the admissable syntypes of A. terrestris Linnaeus is in agreement with the current usage of the name. To reaffirm the traditional usage of this particularly widely used name, a case was made to ICZN by Loken et al. (1994). This sought an Opinion from ICZN (ICZN, 1996) that set aside, by use of its Plenary Power (ICZN, 1985: Articles 78b, 79), the lectotype designation for A. terrestris by Day from application P.H. WILLIAMS of the Code (ICZN, 1985), and then designated a neotype (ICZN, 1996: 64) to conserve the traditional usage of the name for even the narrowest concept of the taxon (ICZN, 1985: Article 75). COMMENT. This species has been introduced into New Zealand (e.g. Gurr, 1957; Macfarlane & Gurr, 1995) (see the comments onB. hortorum, B. ruderatus, and B. subterraneus), Tasmania (Cardale, 1993), and Japan (I. Washitani, in /itt.). It was also apparently introduced into mainland Australia (New South Wales) without persisting (W. Froggatt in Franklin, 1913). Bombus (Bo.) hypocrita Pérez hypocrita Pérez, 1905:30 Bombus (Bo.) patagiatus Nylander patagiatus Nylander, 1848:234 vasilievi Skorikov, 1913:172 Bombus (Bo.) lucorum (Linnaeus) lucorum (Linnaeus, 1761:425 [Apis]), examined 2cryptarum (Fabricius, 1775:379 [Apis]) ?modestus Cresson, 1863:99, not of Eversmann, 1852:134 (= B. modestus Eversmann) ?moderatus Cresson, 1863:109, replacement name for modestus Cresson, 1863:99 monozonus Friese, 1909:674 ?magnus Vogt, 1911:56 2jacobsoni Skorikov, 1912:610, examined ?burjaeticus Kriiger, 1954:277 ?florilegus Panfilov, 1956:1334 ?reinigi Tkalct, 1974b:322, examined @ TAXONOMIC sTaTUS. These bees have received particularly close attention by authors describing the minutiae of colour variation, using at least 186 classi- cal names (see the introduction). At least some of these nominal taxa have been regarded as separate species by some authors (e.g. Rasmont, 1983, 1984, 1988; Scholl & Obrecht, 1983; Scholl et al., 1990; Scholl, Thorp & Obrecht, 1992; Rasmont et al., 1995; Amiet, 1996; Ozbek, 1997; Pamilo et al., 1997). In contrast, B. cryptarum and the North American B. moderatus have recently been treated as conspecific with B. lucorum by Poole (1996). There are conflicting patterns of variation among some characters of these taxa, which are not fully understood (Pekkarinen, 1979; Pamilo ef al., 1984; Amiet, 1996; Pamilo et al., 1997). As far as I can tell from the evidence available at present, separation of the taxa for mapping is still not reliable, at least in some areas of their distribution, and particularly in Central and Eastern Asia, where there appears to be a broad range of variation with some intergradation of character combinations (Williams, 1991). Therefore, CHECKLIST OF BUMBLE BEES because complete mapping of separate taxa is not yet possible for me, B. /ucorum is interpreted here in the broadest sense, to include a complex of similar taxa. However, these taxa require more critical work to clarify population patterns of variation and inherit- ance, even in relatively well known areas such as Britain. O NOMENCLATURE. When Day (1979) came to fix the application of A. terrestris Linnaeus (see the comments on B. terrestris), he had no reason to believe that Linnaeus had not described this taxon from the syntype specimen that was subsequently described as the lectotype (=A. cryptarum Fabricius, see Rasmont, 1988:52, ?= B. lucorum (Linnaeus)). This action brought B. Jucorum (Linnaeus) into subjective junior synonymy with B. terrestris (Linnaeus). To reaffirm the traditional usage of B. terrestris and B. lucorum, a case was made to ICZN by Loken et al. (1994). This sought an Opinion from ICZN (ICZN, 1996) that set aside, by use of its Plenary Power (ICZN, 1985: Articles 78b, 79), the lectotype designa- tion for A. terrestris by Day from application of the Code (ICZN, 1985), and then designated a neotype (ICZN, 1996: 64) to conserve the traditional usage of B. terrestris and B. lucorum (ICZN, 1985: Article 75). COMMENT. This species occurs in Iceland, where it has probably been introduced (Prys-Jones et al., 1981) (see the comments on B. hortorum). Bombus (Bo.) terricola Kirby Terricola Kirby, 1837:273 occidentalis Greene, 1858:12 @ TAXONOMIC sTaTUS. B. terricola and B. occidentalis have been regarded both as conspecific (e.g. Milliron, 1971; Poole, 1996) and as separate species (e.g. Franklin, 1913 [but see p. 239]; Stephen, 1957; Thorp et al., 1983; Scholl et al., 1990). Many specimens from the north west of North America show a reduction in the extent of the yellow bands on gastral terga I and II, with an expansion of the pale pubescence on tergum IV, and so appear to be intermediate or recombinant individuals. Indeed, Stephen’s (1957:74) figure 4 shows several patterns that could represent a continuum of variation between the two forms. Furthermore, Thorp ef al. (1983: fig. 140a) illustrate individuals of ‘B. occidentalis’ from California that look very similar to eastern B. terricola. In view of this, Stephen’s conclusion that there is no intergradation may result from adopting colour crite- ria (identifying B. terricola in the strict sense either by completely black pubescence of female terga V—VI [p. 15] and male tergum IV [p. 19], or by completely yellow pubescence of tergum II [pp. 19, 71], two character states that do not always occur together, even 131 in the east) that could be considered as essentially arbitrary points on a continuum (see the comments on B. fervidus). In view of the existence of apparent intermediates between these nominal taxa in at least part of their range, they are treated here as likely to be conspecific. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Subgenus CULLUMANOBOMBUS Vogt Bombus (Cullumanobombus) Vogt, 1911:57, type-species Apis cullumana Kirby (= Bombus cullumanus (Kirby)) by subsequent designation of Frison, 1927:66 Bremus (Rufocinctobombus) Frison, 1927:78, type-spe- cies Bombus rufocinctus Cresson by monotypy Cullumanibombus Skorikov, 1938a:145, unjustified emen- dation Bombus (Cu.) rufocinctus Cresson rufo-cinctus Cresson, 1863:106 Bombus (Cu.) cullumanus (Kirby) Cullumana (Kirby, 1802:359 [Apis]) examined serrisquama Morawitz, 1888:224 Silantjewi Morawitz, 1892:132 apollineus Skorikovy, 19106:412 @ TAXONOMIC sTATUS. Several of these nominal taxa have been treated as separate species. However, aside from differences in colour pattern, they are closely similar in morphology (Panfilov, 1951). Rasmont (1988) has drawn attention to the co-occur- rence of the white-banded B. apollineus with the yellow-banded B. serrisquama in northern Iran, ap- parently without intermediate individuals. But by analogy, it is possible that this colour difference could be the effect of a single allele for pigment (cf. B. melanopygus, see also the comments on B. keriensis). Until more evidence to the contrary is available from critical studies of patterns of varia- tion, I shall treat them as parts of a single variable species. sy Bombus (Cu.) unicus Morawitz unicus Morawitz, 1883:235 controversus Skarikov, 1910b: 411 @ TAXONOMIC STATUS. B. unicus is similar to B. cullumanus and could possibly be conspecific. How- ever, the male genitalia appear to be more distinct (Panfilov, 1951) from those of the other taxa tradition- ally considered subspecies of B. cullumanus. Bombus (Cu.) semenoviellus Skorikov semenoviellus Skorikoy, 1910b:410 Subgenus OBERTOBOMBUS Reinig Bombus (Obertobombus) Reinig, 1930:107, type-species Bombus oberti Morawitz by monotypy [Obertibombus Skorikoy, 1931:239, incorrect subsequent spelling] Bombus (Obertibombus) Reinig, 1934:167, unjustified emendation @ TAXONOMIC STATUS. Richards (1968) treated Obertobombus as a synonym of the subgenus Sibiricobombus. I have recognised Obertobombus and Sibiricobombus as separate subgenera, because to- gether they do not form a monophyletic group (Williams, 1995). Bombus (Ob.) morawitzi Radoszkowski Morawitzi Radoszkowski, 1876:101, examined hydrophthalmus Morawitz, 1883:240, examined Bombus (Ob.) oberti Morawitz Oberti Morawitz, 1883:238, examined Semenovi Morawitz, 1886:198, examined xionglaris Wang, 1982:432, examined, new synonym duanjiaoris Wang, 1982:444, examined zhadaensis Wang, 1982:444, examined, new synonym @ TAXONOMIC STATUS. B. xionglaris and B. zha- daensis are closely similar to B. oberti in morphology and in colour pattern. These bees occur at high alti- tudes and are not common in collections (Williams, 1991). However, I know of no reason why these P.H. WILLIAMS nominal taxa should not be considered conspecific. Subgenus MELANOBOMBUS Dalla Torre Bombus (Melanobombus) Dalla Torre, 1880:40, type-spe- cies Apis lapidaria Linnaeus (= Bombus lapidarius (Linnaeus)) by subsequent designation of Sandhouse, 1943:569 Bombus (Lapidariobombus) Vogt, 1911:58, type-species Apis lapidaria Linnaeus (= Bombus lapidarius (Linnaeus)) by subsequent designation of Sandhouse, 1943:562 Kozlovibombus Skorikov, 1922a:152, type-species Bombus kozlovi Skorikov, 1910b (=Bombus keriensis Morawitz) in the sense of Skorikov, 1922a (based on males = Bombus pyrosoma Morawitz, a misidentification, see Reinig, 1934:169, requiring designation by ICZN), by subsequent fixation of Sandhouse, 1943:561 Bombus (Kozlowibombus) Bischoff, 1936:10, unjustified emendation Lapidariibombus Skorikov, 1938a:145, unjustified emen- dation ?Bombus (Tanguticobombus) Pittioni, 1939d:201, type- species Bombus tanguticus Morawitz by original designation (provisional synonym) [Bombus (Lapedariobombus )Esmaili & Rastegar, 1974:52, incorrect subsequent spelling] Bombus (MI.) tanguticus Morawitz tanguticus Morawitz, 1886:200 @ TAXONOMIC STATUS. Queens of B. tanguticus are morphologicaly very distinctive (discussed in Williams, 1991), so much so that Pittioni (1939d) considered the species warranted a subgenus of its own. The male remains apparently unknown (the spe- cies occurs at high altitudes in Tibet [= Xizang] and is very rare in collections), so that its precise relation- ships are difficult to resolve at present and a separate subgenus seems premature. Bombus (M1.) simillimus Smith simillimis Smith, 1852b:48, examined [similis Smith, 1854:403, incorrect subsequent spelling] [simillimus Dalla Torre, 1896:548, incorrect subsequent spelling] grossiventris Friese, 1931:303, examined CHECKLIST OF BUMBLE BEES oculatus (Frison, 1933:335 [Bremus]) examined tonsus (Skorikov, 1933b:248 [Sibiricobombus]) examined simillimus Williams, 1991:99, justified emendation Bombus (ML.) richardsiellus (Tkalcii) richardsiellus (Tkalci, 1968a:42 [Pyrobombus]) exam- ined Bombus (ML.) pyrosoma Morawitz pyrosoma Morawitz, 1890:349, examined pyrrhosoma Dalla Torre, 1896:544, unjustified emenda- tion wutaishanensis (Tkalcit, 1968a:39 [Pyrobombus]) exam- ined @ TAXONOMICSTATUS. B. pyrosomahas been con- sidered conspecific with B. friseanus (Bischoff, 1936) and has been considered conspecific with B. formosellus, B. friseanus and B. flavothoracicus (= B. miniatus) (Williams, 1991). From a preliminary analy- sis of colour variation, S.-f. Wang and J. Yao report (in litt.) that these taxa appear to remain discrete and are likely to be separate species. More evidence is awaited. ?Bombus (MI.) formosellus (Frison) formosellus (Frison, 1934:163 [Bremus]) examined @ TAXONOMIC status. B. formosellus has been considered conspecific with B. pyrosoma, B. friseanus and B. flavothoracicus (= B. miniatus) (Williams, 1991), as a disjunct peripheral population on Taiwan. From a preliminary analysis of colour variation, S.-f. Wang and J. Yao report (in /itt.) that these taxa appear to remain discrete and are likely to be separate species. More evidence is awaited. ?Bombus (ML.) friseanus Skorikov friseanus Skorikov, 1933a:62, examined hénei Bischoff, 1936:10, examined @ TAXONOMIC STATUS. _ B. friseanus has been con- sidered conspecific with B. pyrosoma (Bischoff, 1936; Tkalcti, 1961b; Sakagami, 1972) and has been consid- ered conspecific with B. pyrosoma, B. formosellus and B. flavothoracicus (= B. miniatus) (Williams, 1991). From a preliminary analysis of colour variation, S.-f. Wang and J. Yao report (in /itt.) that these taxa appear to remain discrete and are likely to be separate species. More evidence is awaited. ?Bombus (ML.) miniatus Bingham flavothoracicus Bingham, 1897:552, examined, not of Hoffer, 1889:49 (= B. campestris (Panzer)) miniatus Bingham, 1897:553, examined eurythorax Wang, 1982:435, examined, new synonym 133 stenothorax Wang, 1982:439, examined, new synonym @ TAXONOMIC STATUS. B. miniatus has been con- sidered conspecific with B. pyrosoma, B. formosellus and B. friseanus (Williams, 1991). Evidence of intermediates between B. miniatus and B. friseanus is not strong, but not least because so little material is available from where these taxa occur in close proximity in the eastern Himalaya The few workers and males from this area that I have seen are difficult to assign to either taxon with any confidence, although the queens are closer to the colour pattern of B. miniatus (Williams, 1991). More evidence is awaited. B. eurythorax and B. stenothorax are closely similar in morphology and colour pattern to B. miniatus. I know of no reason why these nominal taxa should not be considered conspecific. O NOMENCLATURE. With Psithyrus regarded as being a subgenus of the genus Bombus, B. flavothoracicus Bingham (1897) becomes a junior secondary homonym in Bombus of Psithyrus camp- estris var. flavothoracicus Hoffer (1889) (deemed to be subspecific, see ICZN, 1985: Article 45g(ii)), and therefore the name B. flavothoracicus Bingham is invalid (ICZN, 1985: Article 57c). For this species, the oldest available name is B. miniatus, which becomes the valid name. The only subsequent publications of which I am aware that use the name B. flavothoracicus for this taxon as a species are by Tkalcti (1974b), Wang (1982) and Macior (1990), so this change of valid name is not a serious disruption of common usage. Bombus (M1.) rufofasciatus Smith rufo-fasciatus Smith, 1852b:48, examined Prshewalskyi Morawitz, 1880:342 rufocinctus Morawitz, 1880:343, examined, not of Cresson, 1863:106 (= B. rufocinctus Cresson) chinensis Dalla Torre, 1890{June 25]:139, replacement name for rufocinctus Morawitz, 1880:343; not of Morawitz, 1890[April 30]:352 (= B. chinensis (Morawitz)) waterstoni Richards, 1934:88, examined Bombus (MI.) ladakhensis Richards ladakhensis Richards, 1928a:336, examined, not infrasubspecific after Tkalcti, 1974b:335 phariensis Richards, 1930:642, examined, not infrasubspecific after Tkalcti, 1974b:336 variopictus Skorikov, 1933b:248, examined reticulatus Bischoff, 1936:7, examined Bombus (MI.) semenovianus (Skorikov) semenovianus (Skorikov, 1914a:127 [Lapidariobombus]) examined 134 Bombus (MIL.) incertus Morawitz incertus Morawitz, 1881:229 Bombus (ML.) lapidarius (Linnaeus) Lapidaria (Linnaeus, 1758:579 [Apis]) examined Strenuus (Harris, 1776:131 [Apis]) eriophorus Klug, 1807:265, examined caucasicus Radoszkowski, 1859:482, examined Bombus (ML.) keriensis Morawitz keriensis Morawitz, 1886:199, examined separandus Vogt, 1909:61, examined kohli Vogt, 1909:61, examined, not of Cockerell, 1906:75 (= B. morio (Swederus)) kozlovi Skorikoy, 1910b:413, replacement name for kohli Vogt, 1909:61 tenellus Friese, 1913:86 [alagesianus (Skorikov, 1922a:152 [Lapidariobombus]) published without description] alagesianus Reinig, 1930:89 richardsi Reinig, 1935:341, not of Frison, 1930:6 (= B. rufipes Lepeletier) tibetensis Wang, 1982:439, replacement name forrichardsi Reinig, 1935:341 trilineatus Wang, 1982:441, examined, new synonym @ TAXONOMIC STATUS. Several of these nominal taxa have been treated as separate species (e.g. Skorikov, 1931), although B. keriensis has also long been considered a broadly-distributed and variable species, including both yellow-banded and white- banded individuals throughout much of its range (Reinig, 1935, 1939; Williams, 1991; Fig. 9). B. trilineatus is morphologically closely similar to B. keriensis. | know of no reason why these nominal taxa should not be considered conspecific. Bombus (ML1.) sichelii Radoszkowski Sichelii Radoszkowski, 1859:481, examined [Sicheli Radoszkowski, 1877b:213, incorrect subsequent spelling] tenuifasciatus Vogt, 1909:49, not of Vogt, 1909:49 (= B. pyrenaeus Pérez) after Tkalcii, 1973:266 chinganicus Reinig, 1936:6, not of Reinig, 1936:8 (?= B. bohemicus Seidl) erzurumensis (Ozbek, 1990:209 [Pyrobombus]) examined, new synonym @ TAXONOMIC sTaTUS. Until recently, the white- banded form of B. sichelii has been known from west of the Caspian Sea only from the Caucasus (Reinig, 1935). Now that B. erzurumensis (morphologically closely similar to B. sicheli-* i and with white bands) has been described from Turkey, it could be interpreted as another white-banded, western colour form. By analogy (cf. comments on B. melanopygus), the difference in colour could be the P.H. WILLIAMS effect of a single allele for pigment. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Subgenus SIBIRICOBOMBUS Vogt Bombus (Sibiricobombus) Vogt, 1911:60, type-speciesApis sibirica Fabricius (= Bombus sibiricus (Fabricius)) by subsequent designation of Sandhouse, 1943:599 Sibiricibombus Skorikov, 1938a:145, unjustified emenda- tion [Bombus (Sibericobombus) Kruseman, 1952:101, incor- rect subsequent spelling] Bombus (Sb.) sibiricus (Fabricius) fibirica (Fabricius, 1781:478 [Apis]) examined flaviventris Friese, 1905:514, examined, new synonym ochrobasis Richards, 1930:655, examined, new synonym @ TAXONOMIC sTaTuUS. B. sibiricus and B. flavi- ventris have been regarded as separate species. Females of B. flaviventris are morphologically closely similar to those of B. sibiricus, but differ in having the orange pubescence dorsally between the wing bases and on gastral terga IV—VI replaced with black. S.-f. Wang and J. Yao have kindly shown me the male of B. flaviventris, which 1s closely similar in its genitalia to B. sibiricus. B. ochrobasis appears to differ from B. flaviventris only in the lighter hue of the yellow pubescence of B. ochrobasis. At present I know of no good biological reason why these three nominal taxa should not be regarded as conspecific. More evidence is awaited. COMMENT. B. flaviventris has long been placed in the subgenus Subterraneobombus (e.g. Skorikov, 1922a; Richards, 1930, 1968), although the characters of the females (Williams, 1991) and the males (Wang & Yao, unpublished) agree with the species of the subgenus Sibiricobombus. Bombus (Sb.) obtusus Richards obtusus Richards, 1951:196, examined CHECKLIST OF BUMBLE BEES Bombus (Sb.) asiaticus Morawitz asiatica Morawitz in Fedtschenko, 1875:4, examined longiceps Smith, 1878:8 Regeli Morawitz, 1880:337, examined regelii Dalla Torre, 1896:544, unjustified emendation [miniatocaudatus Vogt, 1909:50, infrasubspecific] miniatocaudatus Vogt, 1911:61, examined, not of Vogt, 1909:56 (= B. soroeensis (Fabricius)) heicens Wang, 1982:430, examined, new synonym huangcens Wang, 1982:430, examined, new synonym flavicollis Wang, 1985:163, examined, new synonym baichengensis Wang, 1985:164, examined, new synonym @ TAXONOMIC STATUS. Several of these nominal taxa have been treated as separate species. B. heicens, B. huangcens, B. flavicollis and B. baichengensis are morphologically closely similar to B. .asiaticus and differ onlyin details of the colour pattern. In the case of the yellow unbanded colour form and the grey banded colour form in Kashmir (Fig. 12), there is evidence of interbreeding, with many recombinant individuals in some localities (Williams, 1991). Aside from differences in colour pattern, these taxa are similar in morphology with a range of variation (Williams, 1991). Until more evidence to the contrary is available from critical studies of patterns of varia- tion, I shall treat them as parts of a single variable species. Bombus (Sb.) niveatus Kriechbaumer niveatus Kriechbaumer, 1870:158 ?vorticosus Gerstaecker, 1872:290, examined (provisional synonym) @ TAXONOMIC STATUS. B. niveatus and B. vorti- cosus have been regarded both as conspecific (Schmiedeknecht, 1883; Handlirsch, 1888; DallaTorre, 1896; Schulz, 1906) and, more recently, as separate species (e.g. Skorikov, 1922a; Pittioni, 1938; Tkalcii, 1969; Reinig, 1981; Rasmont, 1983). As far as I am aware, the white-banded B. niveatus occurs only within the broader distributional bounds of the yellow-banded B. vorticosus (within its “extent of occurrence’ in the sense of Gaston, 1994). Although they differ in the colour of the pale pubescence (Pittioni, 1939a), they are closely similar in morphology (Williams, 1991; Baker, 19965). Pittioni (1938) and Baker (1996b) report that they occur at different alti- tudes, without intermediate colour forms. However, the significance of this is unclear, because Baker (1996b) notes that the white-banded B. niveatus co- occurs with other bumble bees (B apollineus (= B. cullumanus), B. simulatilis (= B. ruderarius)) that also show strong convergences in these areas towards the white-banded colour pattern, while elsewhere they are more broadly distributed in yellow-banded colour forms. By analogy with other species (cf. comments 135 on B. melanopygus, B. keriensis), the difference in colour could be the effect of a single pair of alleles for pigment. It is suspicious that both colour forms show identical variation in the extent of pale fringes to the pubescence on the posterior of tergum ILI. Until more evidence for differences between these nominal taxa other than colour is available from criti- cal studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Sb.) sulfureus Friese sulfureus Friese, 1905:521, examined Subgenus FRATERNOBOMBUS Skorikov Alpigenobombus (Fraternobombus) Skorikov, 1922a:156, type-species Apathus fraternus Smith (= Bombus fraternus (Smith)) by subsequent designation of Frison, 1927:63 Bombus (Fraternobombus) Franklin, 1954:44 Bombus (Fr.) fraternus (Smith) fraternus (Smith, 1854:385 [Apathus]) examined Subgenus CROTCHIIBOMBUS Franklin Bombus (Crotchiibombus) Franklin, 1954:51, type-spe- cies Bombus crotchii Cresson by original designation 136 Bombus (Cr.) crotchiti Cresson Crotchii Cresson, 1878:184 Subgenus ROBUSTOBOMBUS Skorikov Volucellobombus Skorikov, 1922a:149, type-species Bombus volucelloides Gribodo (?= B. melaleucus Handlirsch) by monotypy Alpigenobombus (Robustobombus) Skorikov, 1922a:157, type-species Bombus robustus Smith by subsequent designation of Sandhouse, 1943:597 Bombus (Robustobombus) Richards, 1968:217 COMMENT. Variation within and among the species of this subgenus is particularly poorly understood and a critical review is urgently needed. Bombus (Rb.) melaleucus Handlirsch melaleucus Handlirsch, 1888:228, examined ?volucelloides Gribodo, 1892:119 (provisional synonym) ?vogti Friese, 1903:254 (provisional synonym) ?nigrothoracicus Friese, 1904:188, examined (provisional synonym) melanoleucus Schulz, 1906:267, unjustified emendation @ TAXONOMIC STATUS. Several of these nominal taxa have been treated as separate species. B. volucelloides is closely similar to B. melaleucus, but has been considered to be a separate species (e.g. Milliron, 19735). B. vogti is also closely similar to B. volucelloides, and these two taxa have been consid- ered both as conspecific (e.g. Franklin, 1913; Labougle, 1990) and as separate species (e.g. Milliron, 1973). G. Chavarria (pers. com.) also believes that B. melaleucus, B. volucelloides and B. vogti are all conspecific. In addition, it seems to me that B. nigrothoracicus is more likely to be conspecific with B. vogti than with B. ecuadorius (see the comments on B. ecuadorius). Thus B. melaleucus is interpreted here in a very broad sense, to include much variation that is not yet well understood. Until more evidence to the contrary is available from critical studies of patterns of varia- tion, I shall treat them as parts of a single variable species. P.H. WILLIAMS O NOMENCLATURE. For this species, the oldest available name of which I am aware is B. melaleucus, which becomes the valid name. The name B. volucelloides has been in most common use, although for just part of this species. However, it seems prema- ture to conserve B. volucelloides by suppressing B. melaleucus until the taxa are better understood, be- cause the name B. melaleucus might yet be required for a separate species or subspecies. Bombus (Rb.) ecuadorius Meunier Ecuadorius Meunier, 1890:66 ?butteli Friese, 1903:254, examined (provisional syno- nym) @ TAXONOMIC STATUS. B. butteli is closely simi- lar to B. ecuadorius. They have been considered to be separate species (e.g. Franklin, 1913; Milliron, 1973b), although Franklin conceded that B. butteli (which has grey hairs intermixed on the thoracic dorsum) might be ‘only a variety or subspecies’ of B. ecuadorius (which has the thoracic dorsum en- tirely black). B. ecuadorius females are very rare in collections. For example, Milliron (19736) had seen only five putative specimens (as opposed to 42 specimens of B. butteli). Of these five specimens, four were fe- males, and just one was a male, which is the same specimen as the holotype of B. nigrothoracicus (see the comments on B. melaleucus). This male is la- belled ‘Bolivia / ?Peru’, whereas the rest of Milliron’s B. ecuadorius are from Ecuador, with the exception of one queen from ‘Peru’ (it carries no further locality data). This putative male of B. ecuadorius differs from the females in having yellow hairs intermixed on the front and rear of the thorax. This was not mentioned in the original description of this male (under the name B. nigrothoracicus) by Friese (1904), which Franklin (1913) used subse- quently as the sole basis for associating the male with B. ecuadorius. Currently I favour another possible interpretation. This views the male holotype of B. nigrothoracicus instead as a semi-melanic male of B. melaleucus (the males of B. volucelloides [= B. melaleucus] that I have seen have the thoracic dorsum extensively yellow). This might explain the difference in colour pattern and distribution of this male from other B. ecuadorius. However, a consequence of this inter- pretation would be that the only remaining known difference between B. ecuadorius and B. butteli would be in colour pattern, because the main mor- phological justification for regarding them as separate species (the broader apical process of the gonostylus of the putative male B. ecuadorius, now B. melaleucus in the broad sense) would have been removed. Further evidence is awaited. CHECKLIST OF BUMBLE BEES Bombus (Rb.) robustus Smith robustus Smith, 1854:400, examined ? Bombus (Rb.) hortulanus Friese hortulanus Friese, 1904:188, examined [{hortulans Frison, 1925a:155, incorrect subsequent spell- ing] @ TAXONOMIC STATUS. B. robustus and B. hortu- lanus have been considered both as conspecific (e.g. Franklin, 1913; Frison, 1925a; G. Chavarria, pers. com.) and as separate species (e.g. Milliron, 1973; Asperen de Boer, pers.com.). B. robustus and B. hortulanus are morphologically similar. Among the specimens I have seen, individu- als that have the sides of gastral terga I-II yellow (B. robustus) also have pubescence extending to the mid- dle or almost to the middle of tergum I, and the males have the space between the inner basal process of the gonostylus and the inner apical process narrower than the apical process. Conversely, individuals with the sides of terga I-II black (B. hortulanus) have at least the medial third of tergum I hairless, and the space between the inner processes of the male gono- stylus is wider than the breadth of the apical process. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as separate species. Bombus (Rb.) tucumanus Vachal tucumanus Vachal, 1904:10 Subgenus SEPARATOBOMBUS Frison Bremus (Separatobombus) Frison, 1927:64, type-species Bombus separatus Cresson (= Bombus griseocollis (DeGeer)) by original designation Bombus (Separatobombus) Franklin, 1954:44 Bombus (Sp.) morrisoni Cresson Morrisoni Cresson, 1878:183 Bombus (Sp.) griseocollis (DeGeer) grifeo-collis (DeGeer, 1773:576 [Apis]) separatus Cresson, 1863:165 Subgenus FUNEBRIBOMBUS Skorikov Alpigenobombus (Funebribombus) Skorikoy, 1922a:157, type-species Bombus funebris Smith by monotypy Bombus (Funebribombus) Richards, 1968:214 Bombus (Fn.) funebris Smith funebris Smith, 1854:400, examined Bombus (Fn.) rohweri (Frison) rohweri (Frison, 1925a:144 [Bremus]) @ TAXONOMIC sTATUS. B. funebris and B. rohweri have been regarded both as conspecific (Milliron, 1962) and as separate species (Frison, 1925a; Asperen de Boer, 1993a; G. Chavarria, pers. com.). They have been distinguished with reference to subtle morpho- logical characters as well as to the consistently and strongly differing colour patterns. Both Asperen de Boer (1993a) and G. Chavarria (pers. com.) found that they co-occur at some localities without intermediate colour patterns. Further evidence is awaited. Subgenus BRACHYCEPHALIBOMBUS Williams Bombus (Brachycephalibombus) Williams, 1985b:247, type-species Bombus brachycephalus Handlirsch by original designation 138 @ TAXONOMICSTATUS. B. brachycephalus was not explicitly placed in any subgenus by Richards (1968). I described a separate subgenus Brachycephalibombus for B. brachycephalus and B. haueri (Williams, 19855), in order to maintain monophyletic groups (Williams, 1995). Bombus (Br.) brachycephalus Handlirsch brachycephalus Handlirsch, 1888:244 neotropicus (Frison, 1928:151 [Bremus]) krusemani Asperen de Boer, 1990:1, examined, new syno- nym @ TAXONOMIC STATUS. The description of B. krusemani shows that this nominal taxon, known from a single location, diverges slightly in colour pattern from the otherwise widespread, common and variable Central American species, B. brachycephalus. The information available at present for B. krusemani is consistent with the known range of variation within B. brachycephalus (e.g. Labougle, 1990). Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Br.) haueri Handlirsch Haueri Handlirsch, 1888:234 COMMENT. Franklin (1913) and Labougle (1990) believed that this species is closely related to B. crotchii (although Labougle had not examined any males). Surprisingly, Milliron (1973b) placed B. haueri in his ‘Dentatus-group’, without explanation (B. dentatus is a junior synonym of the Indo-Chinese B. breviceps of the subgenus Alpigenobombus). Possibly Milliron, at least, may have been influenced by Skorikoy (1922a), who placed B. haueri in the subgenus Alpigenobombus (as Alpigenobombus (Alpigenobombus) haueri, which he also listed next toAg. (Ag.) crotchii). However, both sexes of species of the subgenus Alpigenobombus, as it has been accepted recently (Richards, 1968; Williams, 1991), are easily distinguished from any New World bumble bees because they have more teeth on the mandibles. I have examined the morphology of both sexes and, on the basis of cladistic analysis, have grouped B. haueri withB. brachycephalus (Williams, 1985b, 1995) and with B. rubicundus (Williams, 1995). Further evidence is awaited. P.H. WILLIAMS Subgenus RUBICUNDOBOMBUS Skorikov Fervidobombus (Rubicundobombus ) Skorikoy, 1922a:154, type-species Bombus rubicundus Smith by subsequent designation of Sandhouse, 1943:597 Bombus (Rubicundobombus) Richards, 1968:217 Bombus (Rc.) rubicundus Smith [Napensis Spinola in Osculati, 1850:201, published with- out description] rubicundus Smith, 1854:400, examined Subgenus COCCINEOBOMBUS Skorikov Alpigenobombus (Coccineobombus )Skorikovy, 1922a:157, type-species Bombus coccineus Friese by subsequent designation of Sandhouse, 1943:539 Bombus (Coccineobombus) Richards, 1968:214 Bombus (Cc.) coccineus Friese coccineus Friese, 1903:254, examined Bombus (Cc.) baeri Vachal Baeri Vachal, 1904:10 CHECKLIST OF BUMBLE BEES Subgenus DASYBOMBUS Labougle & Ayala Bombus (Dasybombus) Labougle & Ayala, 1985:49, type- species Bombus macgregori Labougle & Ayala by original designation @ TAxonomic status. B. handlirschi was not ex- plicitly placed in any subgenus by Richards (1968), and B. macgregori had yet to be described. I have grouped B. handlirschi with B. macgregori in the subgenus Dasybombus (Williams, 1995). 139 Bombus (Ds.) macgregori Labougle & Ayala macgregori Labougle & Ayala, 1985:50, examined menchuae Asperen de Boer, 1995:47, examined, new syno- nym @ TAXONOMIC STATUS. B. menchuae was de- scribed from a single location and, on the basis of the worker and male I have examined, appears to diverge from B. macgregori only in colour pattern. Until more evidence to the contrary is available from critical studies of patterns of variation, I shall treat them as parts of a single variable species. Bombus (Ds.) handlirschi Friese handlirschi Friese, 1903:255, examined COMMENT. Franklin (1913) believed that this spe- cies is closely related to B. rubicundus. Milliron (1973b) knew ‘of no closely related species in the Western Hemisphere’. 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Valid names are shown in bold. Names in the genus group are shown in capitals. abditus 114 abnormis 124 acutisquameus 104 adventor 107 ADVENTORIBOMBUS 107 affinis 130 afghanus 99 agnatus 126 AGROBOMBUS 107 agrorum 109, 118 alagesianus 134 alboanalis 125 alboniger 128 albopleuralis 115 ALLOPSITHYRUS 102 ALPIGENOBOMBUS 122 alpigenus 122 alpiniformis 120 ALPINOBOMBUS 119 alpinus 120 altaicus 99 americanorum 111 amurensis 121 anachoreta 108 andamanus 128 andreae 125 angustus 123 ANODONTOBOMBUS 123 APATHUS 102 apollineus 131 appositus 122 arcticus 110, 119, 120 ardens 125 arenicola 109 argillaceus 117 armeniacus 105, 118 arvensis 118 ashtoni 104 ASHTONIPSITHYRUS 102 asiaticus 135 assamensis 106 asturiensis 116 atratus 102, 112 atripes 110 ATROCINCTOBOMBUS 128 atrocinctus 129 atropygus 115 audax 130 auricomus 100 avanus 124 avinoviellus 99 azureus 112 baeri 138 baguionensis 125 baichengensis 135 baikalensis 125 balteatus 119 bannitus 107 barbutellus 104 beaticola 125 bellardii \05 bellicosus 112 bicolor 103 bicoloratus 114 bifarius 128 bimaculatus 128 birmanus 106 biroi 126 bischoffiellus 123 bohemicus 104 BOMBELLUS 99 BOMBIAS 100 BOMBUS 99, 129 BOOPOBOMBUS 100 borealis 122 braccatus 106 BRACHYCEPHALIBOMBUS 137 brachycephalus 138 branickii 105 brasiliensis 106, 113 BREMUS 99 breviceps 123 brevivillus 114 brodmannicus 125 bureschi 109 burjaeticus 130 butteli 136 cajennensis 112 calidus 102, 124 californicus 110 caliginosus 127 callophenax 99 campestris 105 canariensis 130 canus 104 carbonarius 112 cardui 118 cascadensis 127 caucasicus 134 celticus 107 centralis 127 CERATOPSITHYRUS 102 charharensis 121 chayaensis 122 chinensis 100, 104, 133 chinganicus 104, 134 chloronotus 105 CHROMOBOMBUS 107 cingulatus 125 cinnameus 107 CITRINOPSITHYRUS 102 citrinus 103, 106 clydensis 119 COCCINEOBOMBUS 138 coccineus 138 cognatus 110 combai 109 CONFUSIBOMBUS 101 confusus 101 consobrinus 115 consultus 103 contiguus 103 controversus 132 convexus 100 coreanus 104 cornutus 104 crawfordi 103 crotchii 136 CROTCHIIBOMBUS 135 cryptarum 130 CULLUMANOBOMBUS 131 cullumanus 131 ezerskii 117 daghestanicus 109 dahlbomii 111 DASYBOMBUS 139 decoomani 104 defector 99 dentatus 123 derhamellus 109 deuteronymus 108 diabolicus 120 difficillimus 121 DIGRESSOBOMBUS 110 digressus 113 diligens 112 distinguendus 121 DIVERSOBOMBUS 115 diversus 115 dolichocephalus 112 duanjiaoris 132 dumoucheli 110 ecuadorius 136 edwardsii 127 elegans 121 elisabethae 105 emiliae 112 EOPSITHYRUS 102 ephippiatus 128 eriophorus 134 erzurumensis 134 eurythorax 133 eversmanni 102, 125 EVERSMANNIBOMBUS 102 eversmanniellus 102 excellens 111 exil 107 exilis 107 EXILOBOMBUS 107 eximius 129 expolitus 104 exul 107 fedtschenkoi 121 ferganicus 105 fernaldae 105 FERNALDAEPSITHYRUS 102 fernaldi 128 ferrugifer 106 FERVIDOBOMBUS 110 fervidus 110 FESTIVOBOMBUS 128 festivus 101, 129 filchnerae 107 flavescens 110, 125 flavicollis 135 flavidus 106 flavifrons 126 flaviventris 134 flavodorsalis 111 flavopilosus 124 flavothoracicus 105, 133 flavus 105, 125 fletcheri 124 florilegus 130 folsomi 128 formosellus 133 CHECKLIST OF BUMBLE BEES formosus 128 fragrans 121 franklini 129 FRATERNOBOMBUS 135 fraternus 135 frigidus \26 friseanus 133 fulvescens 108 fulvofasciatus 107 FUNEBRIBOMBUS 137 funebris 137 funerarius \06 gansuensis 106 genalis |23 genitalis 124 gerstaeckeri 115 gilgitensis 129 gilvus 109 globosus 106 grahami 123 griseocollis 137 grossiventris 132 guatemalensis 103 haematurus 124 haemorrhoidalis 106 handlirschi 139 handlirschianus 100 haueri 138 hedini 104, 109 heicens 135 himalayanus 99 hispanicus 116 hoenei 104, 133 honshuensis 109 HORTOBOMBUS 115 hortorum 1|16 hortulanus 137 huangcens 135 humilis 108 hummeli 115 huntii 128 hydrophthalmus 132 hyperboreus 119 HYPNOROBOMBUS 124 hypnorum 124 hypocrita \30 ignitus 130 imitator 110 impatiens 128 impetuosus 109 incertus 134 indicus 105 inexspectatus 109 infirmus 124 infrequens 124 insularis 103, 124 interruptus 103, 105 intrudens 103 irisanensis 114 Jacobsoni 130 japonicus 130 Jonellus 125 KALLOBOMBUS 118 kashmirensis \22 keriensis 134 kirbiellus 119 kirbyellus 119 klapperichi 104, 125 kohistanensis 124 kohli 112, 134 konakovi 109 Koreanus 117 koropokkrus 124 kotzschi 126 kozlovi 134 KOZLOVIBOMBUS 132 krusemani 138 kuani 106 Kulingensis 114 kurilensis 116 LABORIOPSITHYRUS 102 laboriosus 113 ladakhensis \33 LAESOBOMBUS 106 laesus 106 laevis 107 LAPIDARIOBOMBUS 132 lapidarius 134 LAPPONICOBOMBUS 124 lapponicus |27 latefasciatus 104 lateralis 128 latissimus 129 latofasciatus 99 lefebvrei 118 lemniscatus \24 lepidus 124 LEUCOBOMBUS 129 leucopygus 124 leucurus 124 licenti 104 liepetterseni 107 ligusticus 117 lit 107 linguarius 115 lissonurus 106 longiceps 135 longipes 115 lucorum 130 lugubris 100, 104, 128 luteipes |24 lutescens 109 macgregori \39 maculidorsis 106 maderensis 130 magnus 130 makarjini 99 malaisei 115, 129 margreiteri 99 martensi: 104 marussinus 99 MASTRUCATOBOMBUS 122 mastrucatus 122 maxillosus 104 maxwelli 115 mearnsi 125 medius 113 MEGABOMBUS 115 melaleucus \36 MELANOBOMBUS 132 melanopoda \\7 melanopygus \27 melanurus 121 menchuae 139 MENDACIBOMBUS 99 mendax 99 meridionalis 106, 116 mesomelas 118 METAPSITHYRUS 102 metcalfi 106 mexicanus 114 mimeticus 115 miniatocaudatus 118, 135 miniatus 133 mirus 124 mixtus 104, 127 mlokosievitzii 109 mocsaryi 106 moderatus 130 modestus 114, 125, 130 mongol 121 monozonus 104, 130 montanus 109, 128 monticola 128 montivagus 115 montivolanoides 125 montivolans 107 morawitzi 104, 132 morawitzianus 105 morawitziides 122 morio \12 morrisoni 137 MUCIDOBOMBUS 101 mucidus \02 muscorum \07 mysticus 103 napensis 138 nasutus 123 nemorum 104, 121 neoboreus 120 neotropicus 138 nepalensis 105 NEVADENSIBOMBUS 100 nevadensis 100 niger 112 nigripes 107, 111 nigriventris 112 nigrodorsalis 113 nigrothoracicus 136 nikiforuki 125 nivalis 119 niveatus 135 NOBILIBOMBUS 122 nobilis \22 norvegicus 105 notocastaneus 117 novus \05 nursei 126 nymphae 125 oberti \32 OBERTOBOMBUS 32 obtusus \34 occidentalis 131 oceanicus |25 ochraceus 105 ochrobasis 134 oculatus 133 ODONTOBOMBUS 115 opifex 112 opulentus 108 orichalceus 123 ORIENTALIBOMBUS 106 orientalis 106 ornatus 128 pallidus 107, 111 paradoxus 101 parthenius \24 pascuorum 109 patagiatus 130 pekingensis 117 pensylvanicus \11 151 152 peralpinus 124 perezi 104, 109 pereziellus 107 perniger 117 perplexus 118, 124 persicus \02 personatus 121 phariensis 133 picipes 125 pieli 105 pleuralis 126 POECILOBOMBUS 123 polaris 120 POMOBOMBUS 118 pomorum 118 portchinsky 116 potanini 109 praticola 127 pratorum \25 PRESSIBOMBUS 129 pressus 129 pretiosus 123 priscus 106 proteus 118 prshewalskyi 133 przewalskiellus 116 pseudobaicalensis 109 PSITHYRUS 102 pulcherrimus 122 pullatus 112 pyramideus 104 pyrenaeus 105, 126 pyrenes 99 PYROBOMBUS 123 pyrosoma 133 quadricolor 106 radoszkowskyi 118 redikorzevi 105 regeli 135 reinigi 130 reinigiellus 116 religiosus 118 remotus 109 reticulatus 133 RHODOBOMBUS 118 richardsi 104, 129, 134 richardsiellus 133 roborowskyi 121 ROBUSTOBOMBUS 136 robustus 137 rohweri 137 rotundiceps 125 RUBICUNDOBOMBUS 138 rubicundus 138 rubriventris 112 RUDERARIOBOMBUS 107 ruderarius 109 ruderatus 117 RUFIPEDIBOMBUS 129 rufipes 129 RUFOCINCTOBOMBUS 131 rufocinctus 131, 133 rufocognitus 123 rufofasciatus 133 rufoflavus 125 rupestris 105 saltuarius 116 saltuum 121 sandersoni 126 scandinavicus 128 schrencki 109 securus 118 semenovi 132 semenovianus 1|33 semenoviellus |32 semialbopleuralis 107 senex 114 SENEXIBOMBUS 114 senilis 108 separandus 134 SEPARATOBOMBUS 137 separatus 137 serrisquama 131 shaposhnikovi 100 shillongensis 125 SIBIRICOBOMBUS 134 sibiricus 134 sichelii 134 signifer 124 sikkimi 122 silantjewi 131 simillimus 132 simulatilis 109 simulus 123 sitkensis 127 skorikovi 106 smithianus 110 sololensis 103 sonani 124 sonomae 110 sonorus 111 SOROEENSIBOMBUS 118 soroeensis 118 sporadicus \29 steindachneri 1\13 stenothorax 133 stramineus 122 strenuus 120, 134 subbaicalensis 108 subdistinctus 121 SUBTERRANEOBOMBUS subterraneus 121 subtypicus 124 suckleyi 104 SULCOBOMBUS 101 sulfureus 135 superbus 99 superequester 109 supremus 115 sushkini 116 susterai 104, 105 susteraianus 105 sylvarum 109 sylvestris 106 sylvicola 127 tahanensis 125 tajushanensis 105, 114 TANGUTICOBOMBUS 132 tanguticus 132 tenellus 134 tenuifasciatus 126, 134 terminalis 129, 130 ternarius 128 TERRESTRIBOMBUS 129 P. H. WILLIAMS terrestris 130 terricola 131 tersatus 115 tetrachromus 122, 124 thoracicus 109, 112 THORACOBOMBUS 107 tianschanicus 106 tibetanus 104, 124 tibetensis 134 tichenkoi 115 tonsus 133 transbaicalicus 105 transversalis \12 tricolor 119 TRICORNIBOMBUS 110 tricornis 110 trifasciatus 115 trilineatus 134 trinominatus 114 tristis 108, 119 tucumanus 137 tunicatus 129 turkestanicus 99 turneri 104 UNCOBOMBUS 124 unicolor 104, 109 unicus 132 ussurensis 115 vagans 127 validus 122 vallestris 129 vandykei 127 variabilis 103, 108 variopictus 133 varius 99, 105 vasilievi 130 velox 109 velutinus 112 vestalis 104 veteranus 109 villarricaensis 117 violaceus 112 vogti 136 vogtiellus 109 VOLUCELLOBOMBUS 136 volucelloides 136 vorticosus 135 vosnesenskit 128 waltoni 100 waterstoni 133 weisit 113 wilemani 115 wilmattae 128 wurflenii 122 wutaishanensis 133 xelajuensis 114 xionglaris 132 xizangensis 122 yezoensis 116 yuennanensis 109 yuennanicola 124 yuennanicus 118 zhadaensis 132 zhaosu 127 zonatus 108 Bulletin of The Natural History Museum Entomology Series Earlier Entomology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown, this may also be obtained from the same address. Volume 58 No. | The mealybug genus Planococcus (Homoptera: Pseudococcidae). J.M. Cox. 1989. Pp. 1-78, 40 figs. No. 2 The Simuliidae (Diptera) of the Santiago onchocerciasis focus of Ecuador. A.J. Shelley, M. Arzube & C.A. Couch. 1989. Pp. 79-130, 153 figs (including 2 plates in colour). Volume 59 No. 1 The songs of the western European bush-crickets of the genus Platycleis in relation to their taxonomy (Orthoptera: Tettigoniidae). D.R. Ragge. 1990. Pp. 1-35. A reclassification of the Melanotus group of genera (Coleoptera: Elateridae). C.M.F. von Hayek. 1990. Pp. 37-115. No. 2 The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). S.J. Brooks & P.C. Barnard. 1990. Pp. 117-286. Volume 60 No. | The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). P.H. Williams. 1991. Pp. 1-204. No. 2 Sattleria: a European genus of brachypterous alpine moths (Lepidoptera: Gelechiidae). L.M. Pitkin & K. Sattler. 1991. Pp. 205-241. A review of wing reduction in Lepidoptera. K. Sattler. 1991. Pp. 243-288. Volume 61 No. | Thrips (Thysanoptera) from Pakistan to the Pacific: a review. J.M. Palmer. 1992. Pp. 1-76. No. 2 Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Scaffer and Perissopteryx Warren (Lepidoptera: Geometridae). M. Kruger & M.J. Scoble. 1992. Pp. 77-148. Volume 62 No. | Caloptilia \eaf-miner moths (Gracillariidae) of South-East Asia. Decheng Yuan and Gaden S. Robinson. 1993. Pp. 1-37. No. 2 Neotropical Emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Linda M. Pitkin. 1993. Pp. 39-159. Volume 63 No. | A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: Encyrtidae), parasitoids of the immature stages of economically important insect species (mainly Hemiptera and Lepidoptera). D.W. Huang and J.S. Noyes. 1994. Pp. 1-135. No. 2 A taxonomic review of the common green lacewing genus Chrysoperla (Neuroptera: Chrysopidae). S.J. Brooks. 1994. Pp. 137-210. Volume 64 No. 1 Revision of the neotropical genus Oospila Warren (Lepidoptera: Geometridae) M.A. Cook and M.J. Scoble. 1995. Pp. 1-115. No. 2 Encyrtidae of Costa Rica (Hymenhoptera: Chalcidoidea): the genus Aenasius Walker, parasitoids of mealybugs (Homoptera: Pseudococcidae). J.S. Noyes and H. Ren. 1995. Pp. 117-164. Volume 65 No. | A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae). Y. Zhang and M.D. Webb. 1996. Pp 1-103. No. 2 Encyrtidae (Hymenoptera: Chalcidoidea) of Costa Rica: the genera and species associated with jumping plant-lice (Homoptera: Psylloidea). J.S. Noyes and P. Hanson. 1996. Pp. 105-164. Volume 66 No. | A revised classification of the Asian and Pacific selenocephaline leafhoppers (Homoptera: Cicadellidae). Y. Zhang and M.D. Webb. 1997. Pp 1-121. No. 2 Microtermes in East Africa (Isoptera: Termitidae: Macrotermitinae) S. Bacchus. 1997. Pp. 123-171. Mealybugs of the genera Eumyrmococcus Silvestri and Xenococcus — Silvestri associated with the ant genus Acropyga Roger and a review _ of the subfamily Rhizoecinae (Hemiptera, Coccoidea, Pseudococeidas D.J. Williams Monophyly of the dacetonine tribe-group and its component tribes (Hymenoptera: Formicidae) Barry Bolton An annotated checklist of bumble bees with an analysis of patterns se description (Hymenoptera: Apidae, Bombini) Paul H. Williams ENTOMOLOGY SERIES Vol. 67, No. 1, June 1998