Four New Species of Heteroderidae Including Ekphymatodera n. gen. from California

Four new species and a new genus of Heteroderidae from California are described, and their significance for phylogenetic analysis of the family is discussed. The new genus with type species, Ekphymatodera thomasoni n. gen., n. sp., shares many characteristics with Hylonema Luc, Taylor, &Cadet, 1978, but it is distinguished by its much greater vulva-anus distance and unique cuticular pattern. Hypotheses of relationships of Ekphymatodera and Hylonema with Sarisodera Wouts and Sher, 1971 versus Heterodera Schmidt, 1871 are discussed. Verutus californicus n. sp. is larger than the type species, Verutus volvingentis Esser, 1981, differing in females particularly by the greater distance of its excretory pore from the anterior end. Monophyly of Verutus, which may be an outgroup of all other Heteroderidae, is strengthened by description of V. californicus. Atalodera trilineata n. sp. differs from other ataloderines by having second-stage juveniles with three lateral lines and from the type, Atalodera ucri Wouts and Sher, 1971, by the more subtle cuticular pattern of females and longer juveniles with much longer tails. Atalodera festucae n. sp., with four lateral lines in juveniles, has smaller females than A. trilineata and has a protruding dorsal vulval lip. A unique common ancestor for Atalodera-Sherodera-Thecavermiculatus is supported, and monophyly with Thecavermieulatus andinus Golden, Franco, Jatala, &Astocaza, 1973 is considered.