8 



Concerning the internal phylogeny of Blattaria and of Mantodea, the current ideas are 

 based on the extensive investigations of McKittrick (1964) and McKittrick & Mackerras 

 (1965) (Blattaria) and on the survey in Beier (1968) (Mantodea). In terms of systematics, 

 these authors will be followed in this paper. In some aspects the ideas of these authors 

 are well-founded, but many points are still debatable. 



Beier (1968) divides the Mantodea into 8 families: Chaeteessidae, Metallyticidae, Manto- 

 ididae, Amorphoscelididae, Eremiaphilidae, Hymenopodidae, Mantidae, and Empusidae. 

 These are not grouped into higher-ranked categories. Chaeteessidae are more primitive 

 than all other families in that their hind-wings have a complete second anal-vein and in 

 that their fore-legs are beset with stout setae rather than thorns. In Metallyticidae the 

 second anal vein is vestigial, and in the other families the vein is completely missing. 

 Hence, Chaeteessidae seem to be the first offshoot and Metallyticidae the second. 

 McKittrick (1964) divides the Blattaria into two sister-groups, Blattoidea and Blaberoidea. 

 The Blattoidea, comprising Cryptocercidae and Blattidae, do not reveal a single feature 

 that could be unambiguously regarded as a synapomorphy of the two families. The 

 phylogenetic position of Cryptocercidae has been reanalysed by Grandcolas (1994), who 

 assumes that they are a subgroup of Polyphaginae. However, this assignment is also not 

 very convincing, since many of the homology assumptions upon which this assignment is 

 based are debatable. The Blattidae of McKittrick, comprising Blattinae, Polyzosteriinae, 

 Tryonicinae, and Lamproblattinae, are based on features most of which can be suspected 

 to be plesiomorphic for Blattaria, and the family might be para- or even polyphyletic. The 

 Blaberoidea of McKittrick, including Polyphagidae, Blattellidae, and Blaberidae are 

 founded on the presence of a pair of special compound sclerites in the ovipositor, the 

 crosspieces. However, crosspieces are simple gonangula strictly homologous with those 

 of the other Blattaria (Klass 1995, in press), and the holophyly of Blaberoidea is thus 

 highly questionable. McKittrick's assumption that Blattellidae and Blaberidae are closely 

 related is well-founded. However, the exact relations between the two famihes are 

 uncertain. Interpreting the morphological results of McKittrick (1964) concerning the male 

 and female genitalia from the viewpoint of phylogenetic systematics and parsimony, the 

 Blaberidae would have to be regarded as a rather subordinate subgroup of BlattelHdae; 

 however, not all features are consistently supporting this view. The relations between the 

 various subgroups of Blattellidae, which are Anaplectinae, Plectopterinae, Blattellinae, 

 Ectobiinae, and Nyctiborinae, are also rather unclear. 



The external male genitalia of Blattaria and Mantodea (the phallomere complex composed 

 of the phallomeres) have a highly complicated morphology. The knowledge of these 

 structures is extremely scarce. However, a large potential for phylogenetic research can 

 be supposed to be inherent in them, and this will be used in this paper to contribute to 

 the solution of the basic problems of Blattarian and Mantodean phylogeny. 



The phallomere complex, or at least its anterior part, is concealed within a genital pouch. 

 Abdominal sternite 9 is a saucer-shaped subgenital plate, and the pouch is mainly formed 

 from the interstemal membrane between the stemites 9 and 10, which is deeply invaginated 

 anteriad. The ejaculatory duct opens into this pouch, and the phallomeres are evaginations 

 surrounding the genital opening. The phallomere complex is provided with many sclerites 



