218 
Psyche 
[Vol. 95 
Acroglossal buttons. As noted by Carpenter (1981), Richards 
(1962) incorrectly stated that acroglossal buttons are lacking from 
the ligula of Gayellini. The only species in which they are entirely 
lacking are Paramasaris fuscipennis and cupreus (Fig. 21). These 
structures are also absent from the glossa of Paramasaris brasilien- 
sis, but are present on the paraglossae. This is a transformation 
series in reduction. Other gayellines have the buttons on both the 
glossa and paraglossae (Fig. 22), including Paragayella richardsi, 
the sister-group of Paramasaris, and P. brasiliensis is the sister- 
group of fuscipennis + cupreus (Fig. 2). 
Hypostomal apodemes. These are present in all species, which 
supports the interpretation of synapomorphy with Masarini. They 
are always very narrow (Fig. 23). 
Pronotal carina. Paramasaris and Paragayella are notable for 
having two parallel carinae on the pronotum. One is present at the 
anteroventral margin of the pronotum; the other is posterior to this 
and runs towards the humeri and dorsum of the pronotum (Figs. 
29-31, 40). The second carina shows a transformation series in 
development, ranging from short lateral sections only ( Paragayella , 
Figs. 24, 29), to extending to the dorsum ( Paramasaris brasiliensis, 
Figs. 25, 30), to complete across the dorsum {P. cupreus and 
fuscipennis, Fig. 26). This series apparently corresponds to the 
phylogenetic relationships among these species (Fig. 2). Gayella has 
only the anterior carina (Fig. 7). Euparagiinae also has only the 
anterior carina (Fig. 8), although the humeri are somewhat raised in 
scutellaris. In Masarini the anterior carina is usually blunt, and a 
lateral carina on the humeri may be present (Fig. 37). In all these 
groups, the anterior carina precedes a groove which is frequently 
crenate (secondarily reduced in various Masarini). 
The situation is different in other Vespidae. In Polistinae, the 
structure termed the “pronotal prominence” (Richards, 1978) is 
probably homologous with the anterior carina. Although often 
blunt, it is frequently carinate, and lies at the anteroventral margin 
of the pronotum (Fig. 38). It precedes the pronotal fovea, which is 
sometimes set in a deep depression; there is no lateral groove. In the 
groundplan, there is also a carina on the dorsum (Carpenter, 1989). 
This second carina is usually quite short laterally, and may closely 
approach the anterior carina (Fig. 38). In Polistes the second carina 
extends almost to the ventral pronotal margin in many species, and 
the fovea, which is anterior to this carina, is not preceded by a 
