315 



reduced again in various taxa belonging to subgroup 2.2.3.2.2. However, details of 

 morphology of the Blattella-species are not yet investigated. Thus, there is compelling 

 need for further investigations on the phallomeres of the various subgroups of Blattellidae 

 to resolve these problems in terms of the evolution and polarity of characters. 

 The most parsimonious phylogenetic hypothesis resulting from the discussions in chapter 

 7 is shown in diagram 1 in 7.4. If the species investigated in this paper are true 

 representatives of the Mantodean and Blattarian families and subfamilies they are usually 

 assigned to (compare the systems of McKittrick 1964 and Beier 1968 given in chapter 2), 

 the overall phylogeny of Mantodea and Blattaria is as follows: 



In Mantodea, the basal dichotomy is between Mantoididae and all other families. The 

 second dichotomy is between Chaeteessidae and the remaining families. In Blattaria, the 

 basal dichotomy is between Blattinae + Polyzosteriinae and all other Blattaria. These 

 remaining Blattaria form three groups: The first consists only of the rather isolated 

 Tryonicinae. The second contains Cryptocercidae, Lamproblattinae, and Polyphaginae, the 

 two latter taxa being especially closely related. The third group corresponds to Blattellidae 

 + Blaberidae. Blattellidae are clearly paraphyletic, with Blaberidae being a rather 

 subordinate subgroup. The earliest offshoot within Blattellidae (+ Blaberidae) are the 

 Anaplectinae; the three subsequent offshoots are various taxa previously comprised in 

 Plectopterinae. Blaberidae, Nyctiborinae, Blattellinae, and Ectobiinae form together a 

 holophyletic group. Nyctiborinae and Blaberidae are possibly sister-groups. 

 As regards Blattaria, this phylogenetic hypothesis is in several repects very different from 

 the system of McKittrick (1964): 



- Tryonicinae are not related to Blattinae + Polyzosteriinae. 



- Lamproblattinae are also not related to Blattinae + Polyzosteriinae but to Polyphaginae. 



- Cryptocercidae are not the sister-group of Blattidae but probably of Polyphaginae -i- 

 Lamproblattinae (or possibly of Blattellidae -i- Blaberidae). 



- Blattellidae are paraphyletic since Blaberidae are one of their subgroups. (McKittrick 

 has also expressed this idea in her phylogenetic trees - text figure 3 - but not in her 

 system). 



- Plectopterinae are paraphyletic. 



This hypothesis is based almost exclusively on male postabdominal and genital 

 morphology. Of course, there are still other character complexes which have proved to be 

 useful in analysing Dictyopteran phylogeny, e.g. the morphology of the female genitalia, 

 of the proventriculus (McKittrick 1964), or of the wings. The present knowledge on these 

 character complexes has been revised in a phylogenetic approach in Klass (1995), and a 

 study on the evolution of the ovipositor containing many new results has been completed 

 more recently (Klass, in press). The many characters which are now reliably inteipretable 

 are consistent with the phylogenetic hypothesis presented here. Some characters, however, 

 are still problematic, due to insufficent (in quantity and quality) data. To improve the data 

 base for these character complexes, and also for the male genitalia, by detailed 

 morphological investigations should be the major task of future work on Dictyopteran 

 phylogeny. 



