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J:S:TROD UCTIO:S: :\LH ERI:\LS ;\{ETHODS CONTENTS TRA:S:SP ORT OF NEsTI'.\'. G .\IATERI :\LS MA,IPCLATIO:'\ OF ScE:'\Ts B Y :\hLE Ec GLoss1,1 APID PoLLE'.\'. J-. L,sIPULATIO'.\'.  . The Forelegs ---·----·--·-·  . . Pollen Packing ····---··• •· Typ e I Pollen Packing --· . -·•··--· Typ e II Poll en Packing ·-··-•-·· Ip ~ilateral Type II Poll en Packin g  -· Contralateral Pollen Packing in A pis and Bom b11s --··----···· Contralateral Pollen Packing in l\leliponinae -- The Corbicular Pollen Loa<l in :\Ieliponinae ---· Specializatio ns of Meliponine Pollen Collecting  ····-··· The Euglo ssini -···-·--··-·-·-···-··· E, ·0LuT10-.:ARY Co:-.srnERATIO-.:s ---··· -·-·-···· Ernlution of Poll en Collecting and Tr ansport . ·-·· Ernlution of Corbiculae and their Filling :\!echani sms -··  ----.. Poll en-Packing in the Api<lae  -·--·· DeriYati on of Pollen Manipulati on from Self-Grooming J-.l on:mcnt~ AcK:'\OWLEDGE:\IE'.\'.TS ··  ----···LITERATCRE CITED ·-· ABSTRACT 575 576 577 57S 579 5~0 Sb! Sb! 5il2 5S2 583 5113 587 58S 590 590 593 59--l 59--l 595 597 59S 600 600575 576 577 57S 579 5~0 Sb! Sb! 5il2 5S2 583 5113 587 58S 590 590 593 59--l 59--l 595 597 59S 600 600 Poll en m anipul ation is descri bed for all major groups of Api<lae, and for comparati\'e purpose s, oth er hees are also considered. In prototypic pollen collecting, pollen is remo\' ed from th e fro nt leg s and head, and carried in the crop. In euty pie pollen collecting, it is re m o\'ed from th e fr ont and mid legs as well as surfaces of th e head and thorax and trans ferr ed by th e m id dle legs to the scopa on th e hind legs . Diff eren t derived or metat ypic types of pollen m ani pu lation su pp lement the eutypic beha\'ior and provide for transfer of pollen to th e abd om in al scopa in the ~1egachilidae and from th e abdomen to the hind tibiae in the A p idae. In Ap idae th e bru shy scopa of many other bees is modifie<l into a corbicular scopa, which with th e hin d tibia l sur face fo rms the corbicula. Corbicular filling ( = pollen packing) can be 2 Department of Entomo logy, 3 Department of Systematics an<l Ecology, an<l 4 Snow Entomolo gica l Museum, University of Kansa s, Lawrence, Kansas 660--15, U.S.A . 1 Contribution number I 659 from the Department of Entomology, UniYer~ity of Kansas, Lawrence, Kansas 660--15, U.S.A. 576 T11E UNIVERSITY OF KAN S,\S SCIENCE BPLLETIN achieYecl with little or no moclification of cutypical behaYior; for Apiclae this is called Type I pollen packing, in whic h pollen is put directly into the corbiculae hy the midclle legs. Nesting materials arc packed and tran sportecl in the same way. On the other hand, the corbicula is usually loaded with pollen from its distal end. This is Type II pollen packing. It may be either (a) ipsilateral, a modified eutypic pattern in which pollen is placed on the outside of the tihiotarsal joint by the middle leg of the same sicle, and thenc e pushed basad into the corbicula, or (b) contra lat eral, a deriYed pattern in which pollen transf erred hy the middle legs to th e inner surfaces of the hind basitarsi or swept from the ab<lomen by those basitarsi, is transferred to th e corbicula of the opposite tibia. In pollen-manipulating mo\'ements and structures, certain ~Ieliponinae appear to be the most primiti\'e apids, apparently lacking Type II pollen-packing behavior as well as structures necessary for it. For other l\Ieliponinae, ipsilateral Type II pollen packing is most important, although some have contralateral Type II behavior as part of their repertoire. The small hind hasitarsus without an auricle suggests that contralateral Type II packing is of little importance in most Meliponinae, altho ugh an auricular area can function like the auricle to push pollen basad into the corbicula. Euglossini are inadequately studied, but probably exhibit ipsilateral Type II behaYior. Bombini ancl Apinae are similar to one another in pollenhandling structure and behavior and for both, contralateral Type II pollen packing is a principal method. ?v!odifications of usual pollen-manipulating moYements are seen in scent manipulation by male euglossine bees and in pollen-gleaning actiYity by certain Meliponinae ( Scaura ). Most of th e pollen-manipulation mo\'ements are either the same as self-grooming movements or are opposites of them so that, for example, a structure that is stroked distally for cleaning may be stroked basally to load it with pollen, the cleaning moYement proper serving later for unloading. Presumably such loading movements are deriYed from similar cleaning movements. Only the hind basitarsal moYements that push pollen basad into the corbicula lack counterparts in cleaning or other known behavior.

Pollen manipulation and related activities and structures in bees of the family Apidae