Abstract:
The diversity of the Eriophyoidea is largely unknown and their systematic study mostly entails alpha-taxonomy which is critically important for these mites. Eriophyoid morphology is almost exclusively studied on slide-mounted specimens, and truly permanent specimen slides cannot be prepared and are eventually lost. Shortcomings in taxon descriptions are persistent, and too few morphological characters are available for systematic use, particularly for phylogenetic studies. The fragile, simplified and minute eriophyoid bodies, and the inadequacy of study methods and technology, including preparation and light microscopy, contribute to these problems. The present eriophyoid classification is widely accepted, relatively stable and useful. The major part of the classification, however, is probably artificial, and some taxon delimitations and identifications are becoming increasingly difficult. Scanning electron microscopy (SEM) is only sporadically used to supplement conventional descriptions of eriophyoid mites, and their phylogeny has hardly been studied. In the present study some aspects of eriophyoid systematics and its improvements by incorporating SEM for morphological study and phylogenetic analyses for testing and improving the naturalness of the present eriophyoid classification, are used and appraised. The morphology of about 64 species, mostly from South Africa, was studied with low-temperature (cryo) SEM. The specimens remained turgid and the shape of the mites largely unaltered. A general overview of the contribution of the SEM study towards systematic morphology of the Eriophyoidea is presented. Discrepancies between species descriptions from slide-mounted specimens and the SEM images were found. These include body form, interpretation of structures, resolution and information on minute morphology, and the presence of secretions. Some of these differences were caused by artefacts introduced with slide-mounting of specimens. The SEM study includes a comparative morphological study of the gnathosoma, including a review and appraisal of characters presently used in eriophyoid systematics. New morphological information was found, including new characters that may be of systematic use. Morphology studied with SEM should be routinely incorporated into eriophyoid descriptions, which is not presently the case. The phylogeny of the Eriophyoidea was studied at genus level, using morphological data, to test the monophyly of the present suprageneric taxa. Three data matrices with 66, 60 and 27 informative characters of 316 (including most Diptilomiopus spp.), 64 and 17 eriophyoid ingroup species respectively were analyzed with parsimony analyses, and trees were searched under different parameters. This was done to find different hypotheses regarding the taxon relationships, to roughly assess the robustness of the tree groups, and to use different approaches: a very comprehensive taxon sample, but with low ratio of characters to taxa; an exemplar species sample to improve the ratio between characters to taxa; and a very small taxon sample with a good ratio between characters and taxa, but very little inclusion of variation found in the Eriophyoidea. Most groups found were supported only by homoplasy, but many made biological sense and various potentially monophyletic groups, additional to taxa in the present classification, are proposed for further study. The robustness and convergence of these groups on monophyly are discussed. The Phytoptidae was found to be polyphyletic. Part of the Nalepellinae is probably positioned outside the remainder of the Eriophyoidea, while the rest of the Phytoptidae were positioned in smaller subgroups among the Eriophyidae. The Phytoptinae and Sierraphytoptinae, including Pentasetacus, may group together. The Eriophyidae never grouped together with much support, and the family is both polyphyletic and paraphyletic. The Diptilomiopidae was largely found to be monophyletic, with a relatively strong phylogenetic structure. The Rhyncaphytoptinae is mainly paraphyletic, and the Diptilomiopinae polyphyletic, but part of the Diptilomiopinae may be monophyletic. Three new Diptilomiopus spp. from South Africa are described as part of the study: D. faurius sp. nov. from Faurea rochetiana (A. Rich.) Pic. Serm. (Proteaceae); D. apobrevus sp. nov. and D. apolongus sp. nov. from Apodytes dimidiata E. Mey. ex Arn. (Icacinaceae). They were leaf vagrants not causing any observable symptoms.