6 4 
Psyche 
[March-June 
The least specialized cleaners are found in forms with the least 
specialized and possibly primitive pro-mesothoracic joint. The grades 
of antenna cleaners probably represent sequences of historical develop- 
ment, and the terms primitive, intermediate, and advanced can be 
applied to A, B, and C grade configurations, respectively. 
It has been argued above that two developmental pathways have 
been employed in the parallel differentiation of the carabid antenna 
cleaner, and that the grades may represent historical sequences of 
mechanical improvement. The major cleaning element, the setal 
band, is a modified portion of the horizontal ring of setae about the 
tibial apex. Mechanical improvement of a primitive, i.e., grade A, 
configuration consists of a shift of the setal band from horizontal to 
vertical so that coxal rotation can be used in antenna grooming. 
The pathways are geometric alternatives; the setal band is shifted 
proximally and vertically with or without the posterior spur. Im- 
provement then implies choice of pathway. Polyphyletic adoption 
of a given pathway is readily visualized. Similarity of pathway is 
consistent with, but is not in itself, evidence for phylogenetic rela- 
tionship. Each case must be dealt with separately and evidence 
collected from other functional systems before conclusions are 
reached. When such evidence becomes available, the distributions 
of the several types of antenna cleaner will be germaine to a discus- 
sion of carabid evolution. 
Summary 
1. An antenna cleaner, formed from setal aggregations, is present 
on the protibia of nearly all carabid beetles. 
2. The major cleaning element, the setal band, consists of tightly 
packed bristles arranged in single file. The setal band is serially 
homologous with a section of the apical ring of setae between the 
tibial spurs on the meso- and metalegs. 
3. In the least specialized, and probably primitive antenna 
cleaners, the setal band is apical, horizontal and located between the 
nearly level spur insertions. In others, the band is lengthened, 
developed proximally and may be divided into cleaning and non- 
cleaning sections. 
4. Based on the degree of divergence of the setal band and related 
structures, the forms examined can be divided into three grades. 
5. The most specialized configuration is found in the majority 
of carabid tribes; the least modified antenna cleaner is present in 
only a few tribes. 
