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[Vol. 95 
“prominence” (Fig. 39). Richards (1973) confused the dorsal carina 
in Polistes, behind the fovea, with the anterior carina in other 
polistines, in front of the fovea. Eumeninae, which also have a 
fovea, also have a carina in front of the fovea (Fig. 9), which 
continues across the dorsum in the groundplan (Carpenter and 
Gumming, 1985). The single carina may be a composite structure, 
derived from a state resembling certain polistines with two closely 
approximated carinae (Fig. 38). This is also the case in the ground- 
plan of Vespinae (Carpenter, 1987), where there is a single carina, 
preceding the pronotal fovea and running across the dorsum (Fig. 
10). Stenogastrinae have a highly modified pronotum and lack a 
posterior carina and fovea (Carpenter, 1988), but have a blunt ridge 
anteriorly that may correspond to the anterior carina (Fig. 32). 
Thus, an anterior carina is clearly an ancestral vespid character, 
but considering its diverse form, the posterior carina may have 
evolved multiple times. The alternative interpretation, that it 
evolved once (in the ancestor of all vespids except Euparagiinae), 
requires secondary losses within Gayellini ( Paragayella , Paramasa- 
ris and Gayella independently) and Stenogastrinae. The interpreta- 
tion of nonhomology is more parsimonious, and accords with the 
apparent transformation series in Paramasaris. In any case, the 
separate posterior carina has also been lost on numerous occasions 
within the Masarini and Polistinae (Carpenter, 1989, and in prep.), 
and the possibly composite carina has also been lost several times 
within Eumeninae and Vespinae (Carpenter, 1987; Carpenter and 
Gumming, 1985). Secondary loss also applies to the pronotal fovea, 
present in the groundplan of Eumeninae, Polistinae and Vespinae. 
It has been lost multiple times within Polistinae (Richards, 1978; 
Carpenter, 1989), and probably also in Stenogastrinae. Presently 
available morphological and behavioral evidence supports a sister- 
group relationship between Stenogastrinae and Polistinae + Vespi- 
Figs. 20-28. 20. Paramasaris richardsi, 17X. Lateral view of head. 21-22. Ven- 
tral view of ligula. 21. P. fuscipennis, 25X. 22. Gayella luispenai, 16X. 23. G. 
mutilloides, 14X. Ventral view of head, mouthparts removed. 24-28. Dorsofrontal 
view of pronotum. 24. P. richardsi, 9X. 25. P. brasiliensis, 11X. 26. P. cupreus, 
13X. 27. G. eumenoides S, 10X. 28. G. araucaria 9, 8X. ab: acroglossal buttons; 
ap: anterior pronotal carina; gl: glossa; h: hypostomal apodeme; hp: humeral projec- 
tion; pg: paraglossa; pip: posterior lingual plate; pp: posterior pronotal carina; pr: 
preoccipital carina. 
