10 



The primitively separated proxalaria, axalaria and fulcalaria are. in Neoptera. fused 

 together to form several typical clusters (= strongest neopterous synapomorphy; Kukalovä- 

 Peck 1983. 1991): the humeral plate (HP), the first axillary (lAx). the second axillary 

 (2Ax), the median plate (MED), the third axillary (3Ax), and sometimes the fourth axillary 

 (4 Ax) (not illustrated) (Figs.2a-b, 3) (for additional details see Snodgrass 1935 and 

 Kukalovä-Peck 1983, 1991). In the Coleoptera the basivenalia have fused together to form 

 the first basal plate (IBP) and the second basal plate (2BP) (Fig. 3) (Browne & Scholtz 

 1994). 



The Wing Articulation 



First Axillary (lAx). 



The first axillary is composed of four fused sclerites (Figs. 2b, 4): subcostal fulcalare. 

 subcostal axalare, radial axalare and medial proxalare (FSc + AXSc + AXR + PRM). 

 Recent evidence from Neuroptera clearly shows that the head is composed of only one 

 sclerite (FSc), the neck by two sclerites (AXSc and AXR) and the tail by a single sclerite 

 (PRM) (J. Kukalovä-Peck. personal communication 1993; the interpretation proposed by 

 Kukalovä-Peck & Lawrence (1993:203) that the head is composed of FSc + AXSc. the 

 neck by AXM and the tail by PRM should be replaced by this new interpretation). 

 Proximally lAx articulates with the subcostal proxalare (PRSc) and the radial proxalare 

 (PRR), which are fused together (Fig. 2b). anteriorly with the anterior subcostal basivenale 

 (BScA) (Figs.2b, 3) and distally with the second axillary (2Ax) (Figs.2a, 3). 



Dorsal view (Figs.2a-b. 3. 4): lAx, as mentioned above, articulates with PRSc + PRR 

 proximally (Fig. 2b), BScA anteriorly (Figs. 2b, 3) and 2Ax distally (Figs. 2a. 3). The distal 

 margin of the neck articulates with 2Ax (Figs. 2a. 3). In all Polyphaga this articulation 

 takes the form of a concavity, termed the distal embayment (Fig.4). 



The deltoid-shaped tail is divided into two main sections the proximal arch and the distal 

 arch (Fig.4). In Coleoptera PRR is enlarged posteriorly. The proximal arch of lAx is 

 recurved along the entire length (Fig.4) and articulates just below the distal margin of the 

 more proximal PRR (which is fused to the tergum - Fig. 2b). This junction is termed the 

 PRR articulation. The distal arch articulates with 2Ax (Figs. 2a. 3). Both articulations are 

 mediated by tough membranes. 



Ventral view (Fig.5): Ventrally. the distal margin of the neck folds proximad to form a 

 very broad, crescent-shaped ridge, termed the distal neck ridge. The proximal margins of 

 the head and neck are curved ventrad to form a second, slender ridge, termed the proximal 

 neck ridge. Medially the head and neck are deeply concave. 



The proximal and distal arches of the tail are each often margined by a ridge extending 

 along the entire length of each arch. Often these ridges extend posteriad and fuse to a 

 third ridge which extends along the posterior margin of the tail, forming a triangle with 

 an open top. These ridges are termed the proximal tail ridge, posterior tail ridge and distal 

 tail ridge and are of variable width and length. The ridges and the concavities accom- 

 modate the proximal ridge and lobe of 2Ax. 



