Abstract

The family Tipulidae, comprising the subfamilies Tipulinae, Limoniinae, and Cylindrotominae (3), is the largest family in the Diptera, with 14,000 described species. A very old group of Diptera with many resemblances to the Mecoptera (118, 119), tipulids have a number of derived characters and are perhaps (with the Trichoceridae) the sister group of all other Diptera (56). Tipulids are characterized principally by wing venation, presence of a V-shaped mesonotal suture, deciduous legs, absence of ocelli, and, in larvae, a retractible hemicephalous head capsule. They also differ from other Nematocera in several less obvious characters such as the larval peritrophic membrane (102), sperm structure (5), and cerebral neurosecre­ tory cells (100). Tipulidae probably evolved from ancestors resembling or included in the Upper Jurassic Architipulidae (3), and Limoniinae and Tipulinae were differentiated in the mid-late Paleocene (137). Hennig (56) considers the Limoniinae and Tipulinae to be sister groups; Limoniinae have a distinctive karyotype (134) and aedeagal structure (35). The Cylindrotominae appears to be a relict group that was much better represented in the Tertiary (9); their larvae are distinctive and their tripartite aedeagus appears to represent a primitive condition from which the bipartite limoniine structure and unipartite tipuline condition may have been derived (35). According to Savchenko (119) archaic tipulids probably inhabited sub­ tropical forests, and Rohdendorf (118) states that the modern Tipulidea (Tipulidae and the small families Trichoceridae, Tanyderidae, and Ptychop­ teridae) reach their greatest diversity in the humid tropics. However, crane flies now occur over a very -wide range of latitude, although the distribution of individual species tends to be rather limited; they are common in far

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