319 



area of muscle 110 but does not separate L4d from the L2-sclerotisation within Ive. 

 Character states: (0) absent; (1) present (in Lamproblatta); Polarity: 0>1. 

 Nahublattella and the members of subgroup 2.2.3.2.2. (e.g. Blaberus, Parcoblatta) reveal 

 many apomoiphic features in common, and together they form the certainly holophyletic 

 subgroup 2.2.3.2.: (79)-(89) in 7.4. The morphology near the left posterior end of L2 is 

 in some species of subgroup 2.2.3.2.2. (e.g. Nyctibora) rather similar to Nahublattella and 

 can easily be derived from it, and the division of L2 (articulation AlO) is certainly 

 homologous (fig. 328b, h). However, in all members of subgroup 2.2.3.2.2. the L4d-region, 

 and thus the only element yielding a criterion by which this special division of L2 can be 

 recognised or charcterised, is absent: (95) in 7.4. (In the L2-division as present in subgroup 

 2.2.3.2., L4d has primitively also retained the connection with the common sclerotisation 

 of paa and pda, and this would be a second criterion for recognising the difference to 

 Lamproblatta, whose L2- and L4-division separates L4d from the sclerotisation of paa 

 and pda; fig.329f,g, 6.3.4. However, this criterion can be applied neither to Nahublattella 

 - this species shows a certainly autapomorphic division of sclerite L2E+L4N into a basal 

 and a distal sclerite by the membranous ring 39, fig. 244, which separates L4d from the 

 sclerotisation of paa and pda - nor to subgroup 2.2.3.2.2. for the loss of L4d). 

 Hence, there is a first problem in the L2-division of subgroup 2.2.3.2.2. concerning the 

 assignment of a certain morphology to a certain apomorphic character state: Without 

 having used reciprocal illumination between characters previously, i.e. without having the 

 prehminary assumption of the holophyly of the superordinate subgroup 2.2.3.2. resulting 

 from the consideration of other characters, the characters 1 and 2 would have to be 

 regarded as not assessable in those members of the subordinate subgroup 2.2.3.2.2. which 

 show a L2-di vision (Table la). The assessment and the matrix entries of the characters 1 

 and 2 can only be proper if these two characters are considered in interdependence with 

 other characters having apomorphic states common to Nahublattella and subgroup 

 2.2.3.2.2., i.e. if the probable holophyly of the superordinate subgroup 2.2.3.2. has been 

 recognised previously (Table lb). 



Moreover, some members of subgroup 2.2.3.2.2. reveal a secondary fusion of L2D and 

 L2E+L4N and a secondary loss of muscle 110. That these are reversals results clearly 

 from the hierarchical analysis. Concerning the sclerotisations, the highly apomorphic 

 character state achieved by this reversal conforms exactly with the state present in e.g. 

 Polyphaga (all parts of L2 and L4N form together one sclerite), and it also conforms with 

 the most plesiomorphic state within Blattaria and Mantodea (all parts of L2 and the 

 posterior part of region L41 - L4N not yet differentiated as a separate sclerite - are 

 contained in one sclerite). As regards 110, its absence does likewise correspond with the 

 most plesiomorphic state within Blattaria and Mantodea. 



Hence, there is a second problem in the L2-di vision of subgroup 2.2.3.2.2. concerning the 

 recognition of the polarity and the definition of character states: In elaborating a character 

 state matrix without having used reciprocal illumination between characters previously, 

 the morphology of the sclerotisation and of 110 would have to be regarded as representing 

 rather or most plesiomorphic states of the respective characters (Table la). Only the dis- 

 tribution of the states of other characters and their evidence in terms of phylogeny reveals 



