316 



7.8. Conclusions in terms of the side-reversal of the phallomere complex 



Of the species discussed in this paper, Nahublattella, Supella, Euphyllodromia 

 (Plectopterinae), Blaberus, Byrsotria, Blaptica, and Nauphoeta (Blaberidae) have side- 

 reversed phallomeres, and this is certainly an apomorphic feature. According to Bohn 

 (1987), side-reversal also occurs in some species of Ectobius. All these species belong to 

 subgroup 2.2.3.2. If projected on the phylogenetic tree in 7.4., the distribution pattern of 

 this feature is as follows: 



The three basal offshoots within subgroup 2.2.3.2. give rise to species with side-reversed 

 phallomeres {Nahublattella, Supella, Euphyllodromia). Subgroup 2.2.3.2.2.2.2., the sister- 

 group of Euphyllodromia, contains both normally orientated (Parcoblatta, Nyctibora, part 

 of Ectobius) and side-reversed (Blaberus, Byrsotria, Blaptica, Nauphoeta, part of Ectobius) 

 species. (Ectobius can be assigned to this subgroup since the females show the advanced 

 rotation of the oothecae, (117) in 7.4.). 



This distribution can be interpreted in two ways: (1) Side-reversal is a ground-plan feature 

 of subgroup 2.2.3.2. Nahublattella, Supella, Euphyllodromia, the Blaberidae, and the 

 respective species of Ectobius have retained this orientation. Parcoblatta, Nyctibora, and 

 the other species of Ectobius have achieved their normal orientation by a second side- 

 reversal (independently in the various taxa concerned). (2) In the ground-plan of subgroup 

 2.2.3.2. the phallomere complex is still normally orientated. The basal offshoots 

 Nahublattella, Supella, and Euphyllodromia, and also the Blaberidae and the respective 

 species of Ectobius have independently reversed the phallomere complex. 

 Alternative (1) is highly supported by the fact that the three basal offshoots of subgroup 

 2.2.3.2. are side-reversed. However, a definitive decision, whether (1) or (2) or any 

 combination of these possibiHties is true, is not possible at the present state of knowledge, 

 and more species will have to be investigated. At least, it is strongly suggested that the 

 orientation of the phallomere complex, side-reversed or normal, is not a very good criterion 

 for phylogenetic conclusions. 



7.9. Remarks on the procedure in the phylogenetic analysis and on character lists 

 and character state matrices 



Character lists, describing the characters, their states, and the assumed polarities, and 

 character state matrices, describing the distribution of the character states over the taxa, 

 have the function to present all the character states used and their distribution 

 independently of any previous assumptions on phylogeny - as an objective basis for the 

 phylogenetic analysis or as a starting-point for a computer-based cladistic analysis. The 

 applicability of this method of presentation in the frame of an analysis concerned with a 

 very complex type of character evolution, as it has been found in the male genitalia of 

 Blattaria and Mantodea, is discussed here. 



The procedure in the present phylogenetic analysis is mainly hierarchical: Mantodea and 

 Blattaria have been, in the frame of the species investigated, regarded as sister-groups 

 (Isoptera disregarded); this basic assumption is well-founded (Hennig 1969, Klass 1995). 



