2) Arista bare (0), sparsely and short pilose (1), or plumose (densely and long pilose)(2). The aristal pilosity varies greatly in both Graptomyza and Copestylum. In Graptomyza most species have the arista sparsely pilose with short hairs, which I accept as the ground plan condition (as inclusa), some species have only a few very short aristal hairs, one undescribed species from Australia has no aristal hairs, and some species have long and numerous hairs (as in longirostris). In Copestylum, there are a few species groups with distinctive aristal pilosity, but these are all clearly derived from the basic plumose condition. One species, pseudotachina, has the arista bare, a secondary reduction.

3) Mesoanepisternum bare (0) or pilose (1).

4) Mesokatepisternum continuously pilose (0) or bare medially (1).

5) Mesoanepimeron bare posteriorly (0) or pilose (1).

6) Mesokatepimeron (barrette) bare (0) or pilose (1).

7) Postalar wall bare (0) or pilose posteriorly (1).

8) Scutellum without (0) or with preapical depression (1).

9) Medius with apical portion (apical crossvein) processive (0), arcuate and strongly recurrent (1) or straight (2).

10) Cell r4+5 (apical cell) open (0), petiolate (1) or bulbous apically (2). Within Copestylum, the apical cell varies from widely open, to petiolate and bulbous. I accept the widely open state as the ground plan condition for Copestylum.
11) m2 present (0) or absent(1).

12) Larvae saprophagous (0) or specialized inquilines in nests of social Hymenoptera (1).

The taxa examined were:

Copestylum (apicalis Loew, compactus Curran, fornax Townsend, gibbera Schiner, hirtipes Macquart, marginatum Say, mexicanum Macquart, trituberculatum Thompson, tympanitis Fabricius);
Ferdinandea (cupreus Scopoli);
Graptomyza (alabeta Mutin, doddi Ferguson, flavicollis Ferguson, inclusa Walker, liberia Greene, longirostris Wiedemann, maculipennis de Meijere, microdon Osten Sacken, nigripes Brunetti, plumifer Ferguson, signata Walker);
Ornidia (all species);
Volucella (bombylans Linnaeus, decolorata Walker, elegans Loew, inanis Linnaeus, inflata Fabricius, jeddona Bigot, linearis Walker, nigricans Coquillett, pellucens Linnaeus, rotundata Edwards, tabanoides Motschulsky, trifasciata Wiedemann, and zonaria Poda).

I now accept the diphyletic origin of the Old World volucellines as Graptomyza are now known to be saprophagous, not specialized inquilines. Also, Tachinosyrphus is no longer accepted as a genus distinct from Copestylum (new synonymy) as that arrangement left Copestylum as undefined. The above cladistic formula, which is the same as my 1972 phylogeny, is derived when the type species are used as exemplars (CI = 87, RI = 71, 1 tree (Hennig86 used)). The possibility that Ornidia is a specialized subgroup of Copestylum rests on the evaluation of one character, the pilosity of the posterior portion of the anepimeron. In my previous analysis, I considered the pilose condition to be primitive in all situations. Hence, Copestylum was defined by a bare posterior anepimeron. However, under a strict outgroup criterion, the bare condition is primitive within the tribe Volucellini. Copestylum is a large genus (350+ species) with many distinct groups. When exemplars of these groups are added to the analysis and strict outgroup criterion is used, a far different statement of relationship results (Volucellini = Graptomyza + (Volucella + Ornidia + Copestylum), consensus of some 100 trees). Thus, until Copestylum can be revised, the relationships of Ornidia will remain uncertain.

Characters of Volucelline Taxa Taxa Characters