A three-dimensional model of the stomatostylet and associated structures has been reconstructed from serial thin sections of Aphelenchus avenae, a representative of Tylenchomorpha, a group including most plant parasitic nematodes. The reconstruction is compared with previous work on bacteriovorous cephalobids and rhabditids to better understand the evolution of the stylet and its associated cells. Two arcade syncytia ("guide ring") line the stylet shaft, supporting the hypothesis that the stylet shaft and cone (into which the shaft extends and which is not lined by syncytia) are homologous with the gymnostom of cephalobids, the sister taxon of tylenchids. Epidermal syncytia, HypA, HypB, HypC, and HypE, line the cephalic framework, vestibule, and vestibule extension, congruent with the hypothesis that these components are homologous with the cephalobid cheilostom. Relative to outgroups, HypC is expanded in A. avenae, enclosing sensilla that fill most of the cephalic framework. The homolog of syncytium HypD in the cephalobid Acrobeles complexus is not observed in A. avenae. Arcade syncytia are reduced compared with those of cephalobids. Stylet protractor muscles in A. avenae are homologous with the most anterior set of radial muscles of cephalobids. Observations to date test and verify our previous hypotheses of homology of the stomatostylet with respect to the stoma of bacteriovorous outgroups. Reconstruction of the stegostom and pharynx will provide further tests of homology and evolution of feeding structure adaptations for plant parasitism.
KeywordsAcrobeles complexus; arcade syncytia; Caenorhabditis elegans; Cephalobomorpha; feeding; fine structure; free living; homology; modeling; muscle; phylogeny; stoma; syncytium; transmission electron microscopy Molecular phylogenies often raise questions of morphological evolution where they impose a new paradigm on classical scenarios reliant on opposing ideas of phylogeny. One pressing case of incongruence has been shown in nematodes regarding the evolution of the Tylenchomorpha, which include most species of plant parasitic nematodes, as well as many insect parasites and *Correspondence to: Erik J. Ragsdale, Department of Nematology, University of California, Riverside, CA 92521. erik.ragsdale@email.ucr.edu. Additional Supporting Information may be found in the online version of this article.
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NIH-PA Author ManuscriptNIH-PA Author Manuscript NIH-PA Author Manuscript fungivores. With finer phylogenetic resolution based on molecular analyses, it is apparent that evolution of feeding structures for a parasitic lifestyle can no longer be explained by traditional scenarios, with which new hypotheses of relationships are incongruent. Accumulating evidence suggests that Tylenchomorpha, with a specialized stomatostylet, are sister taxon to or nested within the Cephalobomorpha (Aleshin et al., 1998;Blaxter et al., 1998;Holterman et al., 2006;Meldal et al., 2007;Nadler et al., 2006;Smythe and Nadler, 2006), which lack a stylet and instead have an open...