Tolhuaca Nepaloptila Matrioptila Tolhuaca Nepaloptila Matrioptila Tolhuaca Tolhuaca Tolhuaca Protoptilinae The Neotropical caddisfly genus To l h u a c a (Trichoptera: Glossosomatidae) Robertson, Desiree R. Holzenthal, Ralph W. Zootaxa 2005 1063 53 68 7XGZ [404,517,358,384] Insecta Glossosomatidae Tolhuaca Animalia Trichoptera 2 55 Arthropoda genus   Schmid (1964)originally placed  Tolhuacain the family Sericostomatidae, commenting on the similarity of the bifid tergum X of the male genitalia to that of Brachycentridae, at that time a subfamily within Sericostomatidae. Flint (1967)later transferred  Tolhuacato Protoptilinaeupon discovery that the wing figures in the original description were mislabelled with those of  Austrocentrus griseus Schmid 1964(Helicophidae). Since that time, there has been nothing published regarding the placement of  Tolhuacain relation to the other protoptiline genera. When Schmid (1964)first described the genus he wrote, “Dans l’état actuel de nos connaissances, il est impossible d’assigner une position phylétique au genre  Tolhuaca, dont la nervulation assez complète, surtout aux ailes antérieures, contraste avec l’extrême simplification et spécialisation des génitalia…” [“In the current state of our knowledge, it is impossible to assign a phyletic position to the genus  Tolhuaca, whose rather complete venation, especially in the forewings, contrasts with the extreme simplification and specialization of the genitalia…” Translation from Schmid (1964)]. The male genitalia ( Figs. 5A, 7A) have completely lost both the inferior and preanal appendages, and sternum IX has been reduced to nothing more than a thin, ventral strap. The phallic apparatus ( Figs. 5A, 5D, 5F, 7A, 7D) consists of a greatly enlarged, but simple tubular phallobase, and an eversible membranous endotheca. These apparently derived conditions of the male genitalia contrast with other features that imply a basal placement of  Tolhuacawithin Protoptilinae. Published works regarding protoptiline phylogeny are few and far between. Morse and Yang (1993)discussed some genera from the East Palaearctic and Oriental regions, and also provided a very useful table comparing wing venation among 15 recognized genera. Ross (1956)provided an early discussion of possible phylogenetic relationships among protoptiline genera and considered the genus  Matrioptila Ross 1956to be most primitive based on characteristics of the male genitalia and wing venation. In  Matrioptila, segment IX is complete and ringlike, tergum X consists of a pair of simple lobes, and inferior appendages are present. Forewing veins Cu1 and Cu2 are often fused in Protoptilinae, but in  Matrioptila, they are separate and distinct along their entire lengths and in the hind wing, Cu1 is branched. These genitalic and venational characters are primitive according to Ross (1956). Kimmins (1964)described a new genus,  Nepaloptila, from Nepaland placed it in Protoptilinae. The genus shares some characteristics of the male genitalia with  Matrioptila, but Kimmins (1964)considered  Nepaloptilato be more primitive, based on its retention of apical fork V in the forewing, which is usually absent in other Protoptilinae. The venation of  Tolhuaca( Figs. 3, 4) is quite similar to that of  Nepaloptila( Kimmins 1964), differing only slightly. In  Tolhuaca, Scis distinct from R 1 inthe forewing, whereas in  Nepaloptila, Scand R1 are fused near the wing margin. The 2 genera also differ in the position of crossveins in the forewing: those of  Tolhuacaform a relatively straight transverse cord along the anastomosis whereas in  Nepaloptila, they do not. In the hind wing, A2 is present in  Tolhuaca, but absent in  Nepaloptila.  Although  Tolhuacawould seem to have more specialized male genitalia than  Nepaloptilaand  Matrioptila, other characters suggest  Tolhuacais the more primitive genus. In  Nepaloptilaand  Matrioptila, the foretibial spur has been lost, whereas in  Tolhuaca, although reduced considerably, the spur is retained (Fig. 2). The genus  Tolhuacaalso has two small setal warts on the mesoscutellum ( Fig. 1), which Ross (1956)considered to be primitive; these warts are absent in all other known protoptilines. The female genitalia ( Figs. 6, 8) consist of an elongate oviscapt and 2 pairs of long, rod­like apodemes on segments VIII and IX. The presence of these apodemes is pleisiomorphic within Amphiesmenoptera ( Kristensen 1984). The retention of the foretibial spur, presence of mesoscutellar setal warts, and the structure of the female genitalia suggest that  Tolhuacadeserves a basal placement within the subfamily Protoptilinae.