Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al (1988): phylogenetic systematics and revised classification.

Taxonomic schemes for the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al., (1988) have been unstable due to the large number of genera and the paucity of known reliable characters. Reliable characters are essential when using phylogenetic inference in developing a natural classification. Morphological and developmental studies using light, scanning and transmission electron microscopy have revealed the new characters of host response, en face patterns, phasmid structure and female cuticular layers. These techniques also gave us insight into the homoplasy and polarity of many characters, revealed previously undetected character states and clarified misinterpreted character states. A matrix with the 19 most reliable characters is proposed for 20 operational taxonomic units (OTUs) and we employ this matrix for comparing computer generated phylogenetic analyses of the PHYLIP and PAUP packages. PAUP was deemed the more reliable parsimony algorithm for phylogenetic analysis of the Heteroderinae (Fink, 1986; Platnick, 1987). Monophyly of Atalodera + Sherodera + Thecavermiculatus (tribe Ataloderini), and Cactodera + Heterodera + Afenestrata, as well as Punctodera + Globodera + Dolichodera is supported by both programs. Most importantly, analyses strongly support monophyly of all cyst-forming genera (tribe Heteroderini) contrary to previous hypotheses of repeated evolution of the cyst (Wouts, 1985). In addition, monophyly of the Heteroderini with the Ataloderini is demonstrated. PAUP indicates monophyly of Sarisodera + Rhizonema + Bellodera + Hylonema and Ekphymatodera (tribe Sarisoderini new rank). Monophyly of the Sarisoderini was at first only weakly supported, but, subsequently, the reduced width of the submedial lips of second stage juveniles and males was recognized as a synapomorphy which strengthened subsequent PAUP trees and monophyly of the tribe. The present study rejects as paraphyletic or polyphyletic several previously proposed combinations, including Thecavermiculatus sequoiae (versus Rhizonema sequoiae), Sarisodera africana (versus Afenestrata africana), Dolichodera andinus (versus Thecavermiculatus andinus). The question whether T. andinus is a distinct genus, was not resolved due to insufficient data. PAUP supports our previous observations that Cactodera betulae is intermediate in a transformation series between other Cactodera and Heterodera: it also indicates these species as bring monophyletic with Heterodera + Afenestrata, but not with other Cactodera. Although these phylogenetic analyses strongly support some relationships, they indicate unresolved alternative hypotheses for others. Meloidodera (tribe Meloidoderini) and Cryphodera (tribe Cryphoderini) must be investigated for consideration of a possible synapomorphy not included in the present data matrix. Future studies are proposed to more clearly define the monophyly of the Heteroderini, as well as the Sarisoderini. Tests are also proposed to clarify questions of the monophyly of Verutus (tribe Verutini new rank) with the Heteroderinae versus other Tylenchida.