232 



Thus, the outgroup comparison with Ensifera as well as biological properties of Mantodea 

 suggest that the copulation procedure of Blattaria is plesiomorphic and that of Mantodea 

 apomorphic: Phase (1), in which Blattaria make use of their hla-hook, can be regarded as 

 secondarily lost in Mantodea. Additionally, since the phallomeres of Mantodea and e.g. 

 Periplaneta are rather similar in their morphology (and completely different from those 

 of Ensifera), it might be assumed that the way the Mantodean phallomeres functioned 

 before the copulation procedure has changed was similar to that of the Blattarian 

 phallomeres, and that a hla-hook was present for initial seizing. Though these ideas are 

 highly speculative, it is at least plausible that hla and the associated elements L3 and 114 

 were present in the common ground-plan of Blattaria and Mantodea and have been lost 

 in Mantodea. The same might also be true of the nla-process, which is present in many 

 Blattaria (fig.69, 98, 212) but never in Mantodea. nla probably has the function to stiffen 

 the sclerotisation at and near the 114-insertion, and if hla and 114 are lost an additional 

 loss of nla could be expected. 



6.4.3. Homology relations and character states of the elements in Blattaria 



The hla-hook is present in all Blattaria. The homology of all these hla is suggested by 

 their position in the leftmost part of the left complex, by their similar shape, and by the 

 presence of a special sclerite L3 occupying the distal part of hla (L3, however, can be 

 very different in its extension). Apart from these superficial features, additional similarities 

 between certain species confirm this homology assumption. The most important question 

 in this context is whether the main muscles of the hla-hooks (called 114 in most species) 

 are homologous. 



Archiblatta, Periplaneta, and Eurycotis 



The homology of L3, hla, and 114 of these species is quite evident. (1) The hla-base takes 

 the same relative position: right-ventral to the L41-region, left-posterior to the L4n-region 

 with the nla-process, and left-anterior to sclerite L4F (fig. 54, 56, 66, 67). (2) L3 occupies 

 the entire hla except for the basalmost part (30 in fig.65-67). (3) The tip of hla is two- 

 pointed (fig. 53, 65). (4) In Periplaneta and Eurycotis the main muscle of hla (114c in 

 fig. 72) comes from the L4n-region on and near the nla-process and inserts immediately 

 behind si (fig. 70). However, only Eurycotis has one accessory hla-muscle 114d (fig. 73) 

 - possibly a subdivision of 114c. 



Cryptocercus and Lamproblatta 



The hla-base has a similar position relative to the insertions of 12 and si (fig. 156, 157, 

 184, 185) as in Eurycotis (fig.70), and the anterior insertion of the main muscle of hla 

 (114 in fig. 157, 184, 185) is likewise immediately behind the sl-insertion (fig. 157, 158, 

 184, 185). Thus, homology can be assumed for the hla, L3, and 114 of these three species 

 (homology discussion of si in 6.9.). Cryptocercus has one accessory hla-muscle 119 

 (fig. 156); Lamproblatta has two, 122 and 123 (fig. 184- 186). These accessory muscles and 

 114d of Eurycotis all have different insertions, and homology relations are not assumed. 

 In Cryptocercus - as compared with the previous species, Polyphaga, and Ergaula (see 

 below) - the base of hla is more posteriorly, and hla is shorter (fig. 151). 



