In a study of relationships among selected cyst-forming and noncyst-forming species of Heteroderoidea, combined sequences comprised of DNA from part of the conserved 18S ribosomal RNA gene (rDNA) plus the complete ITS rDNA segment were more similar to analyses based on the ITS data alone than to analyses based on the 18S data alone. One of the two noncyst-forming species, Ekphymatodera thomasoni, grouped with cyst-forming species of Heteroderoidea. Bilobodera flexa, also a noncyst-forming species, was separated from all the other taxa by a long branch. Afenestrata koreana, with a weakly sclerotized cyst, grouped closely with H. bifenestra. These observations suggest that phylogenetic analyses using molecular data may aid in our understanding of the evolution of cyst formation in nematodes, including the possibility of secondary loss. The usefulness of molecular phylogenetic analyses in nematodes may depend more on the particular selection of taxa than on mere addition of data from additional genes.
Afenestrata koreana; Bilobodera flexa; Cactodera betulae; Ekphymatodera thomasoni; Globodera; Heterodera bifenestra; ITS1; ITS2; nematode; phylogenetic analysis; ribosomal DNA; 5.8S gene; 18S gene
Fine structure of the stoma, including the cheilostom, gymnostom, and stegostom of Bunonema sp. and Teratorhabditis palmarum was compared with Caenorhabditis elegans to consider fine structural characters that may be phylogenetically informative. The stegostom, enclosed by the anterior end of the pharynx, includes a triradiate lumen surrounded by radial cells (interradial or pairs of adradial cells) repeated in the dorsal and subventral sectors; in Rhabditina, typically the stegostom includes anteriorly two sets of epithelial and posteriorly two sets of muscular radial cells. These muscle cells are anteriorly m1 and posteriorly m2. In Bunonema sp., unlike T. palmarum and C. elegans, the stegostom has a third set of interradial epithelial cells. In Bunonema sp., m1 is expressed by three interradial cells, whereas in T. palmarum and C. elegans m1 is three pairs of adradial muscle cells (i.e., six cells). In all three taxa m2 is expressed as three pairs of adradial muscle cells. Posterior processes of adjacent adradial cells fuse, and closely apposed nuclei may present a figure-eight shape. However, in Bunonema the three interradial m1 cells each have a long posterior process enclosing two separate round nuclei. In combination with additional characters, these diverse stoma features may prove phylogenetically informative. Specifically, the radial epithelial cells of the stegostom appear to be a synapomorphy consistent with a bunonemid-diplogastrid-rhabditid clade, whereas a thickening in the dorsal sector of the stoma cuticle lining is interpreted as a synapomorphy supporting a bunonemid-diplogastrid clade.
Bunonema sp.; Caenorhabditis elegans; cell fusion; DAPI; Diplogastrina; fine structure; nuclei; SEM; TEM; Teratorhabditis palmarum
The ultrastructure of the body wall cuticle in Acrobeles complexus, Cervidellus alutus, and Zeldia punctata was studied as a step toward understanding biological diversity within Cephalobinae, and to discover new characters for phylogeny-based classification of the suborder. In each species the cuticle consists of cortical, median, and basal layers. The cortical layer includes an external trilaminate and internal granular zone; the basal layer is striated. In Z. punctata the median layer is electron-lucent, vacuolar, and penetrates the cortical layer; it also includes periodically dense columns that apparently correspond to punctuations visible with light microscopy. In contrast, the median layer of the body wall cuticle in A. complexus and C. alutus is bisected by a zone that undulates parallel to the nematode surface and with periodicity corresponding to annuli. Phylogenetic analysis, using derived cuticle patterns of Cephalobinae, requires an understanding of ecological pressures that could result in convergent evolution of cuticle characters.
Acrobeles complexus; Cephalobidae; Cervidellus alutus; cuticle; nematode; phylogeny; ultrastructure; Zeldia punctata
Three new species of Nothacrobeles are described from localities in the Mojave Desert, southern California. Nothacrobeles triniglarus n. sp. is characterized by the presence of a long post-vulval sac and three tubular adoral projections. Both N. spatulatus n. sp. and N. nanocorpus n. sp. are smaller than any other known species within the genus. Nothacrobeles spatulatus n. sp. has labial probolae that are short and spatulate without a basal ridge, whereas those of N. nanocorpus n. sp. are flattened and plate-like. Furthermore, N. nanocorpus n. sp. is unique by its extremely short esophageal corpus (less than 25 µm long in adult females) and the small size of its guard processes. An emended diagnosis of the genus is given to accommodate distinctive characteristics of these new species. A table comparing the 11 valid species of Nothacrobeles is presented.
Cephalobidae; morphology; new species; Nothacrobeles nanocorpus n. sp.; Nothacrobeles spatulatus n. sp.; Nothacrobeles triniglarus n. sp.; SEM; southern California; taxonomy
The ultrastructure of the isthmus and basal bulb (postcorpus) of Diplenteron sp. (Diplogasterida) was revealed through transmission electron micrographs of serial sections. The postcorpus is glandular and muscular. There are 26 cells in the postcorpus, including 6 marginal (two sets of three), 6 muscle (two sets of three), 3 gland, and 11 nerve cells. Most of the cell bodies, including the nuclei, are in the basal bulb. Unlike Caenorhabditis elegans, Diplenteron sp. has three gland cells. The glands are embedded in a muscular framework in both taxa, but each gland cell is much bigger in Diplenteron sp. than in C. elegans. Each of the anterior set of three marginal cells is located at the apex of the esophageal lumen and overlaps slightly with one of the posterior sets of three marginal cells. All six marginal cells in Diplenteron sp. have homologs in C. elegans. The anterior set of radial muscle cells is V-shaped and is homologous to m5 muscle cells in C. elegans. The posterior set of muscle cells appears to be homologous to m6 muscle cells in C. elegans. Diplenteron sp. does not have muscle cells corresponding to the m7 cells associated with the "grinder" in C. elegans, which is absent in diplogasterids. The single saucer-shaped muscle cell, m8, covering the posterior wall of the basal bulb in C. elegans was not observed in Diplenteron sp. The structure of the esophageal-intestinal junction in Diplenteron sp. is similar to that of C. elegans in being composed of five epithelial cells. Neurons appear to be more abundant in Diplenteron sp. than in C. elegans. Ultrastructure of the esophagus in diplogasterids, rhabditids, cephalobids, and tylenchids will be useful in testing classical and recent competing hypotheses of secernentean phylogeny.
basal bulb; Caenorhabditis elegans; Cephalobina; Diplenteron; Diplogasterida; esophagus; homology; nematode; postcorpus; Rhabditida; transmission electron microscopy; ultrastructure
Electron and light microscopy were used to study the dorsal gland (DG) and the two subventral glands (SvG) of seven developmental phases of Nacobbus aberrans: pre-parasitic second-stage juveniles (J2), parasitic J2, third- (J3) and fourth- (J4) stages, migratory females, young sedentary females, and mature sedentary females. In each developmental phase the level of esophageal gland activity, was estimated by the abundance of organelles associated with secretory pathways, including endoplasmic reticulum, ribosomes, Golgi, multivesicular bodies, and secretory granules. All esophageal glands were metabolically active in all J2 examined, although only in parasitic J2 were there numerous secretory granules in the esophageal gland extensions and ampullae. No evidence of secretory activity was observed in the esophageal glands of the coiled and relatively inactive J3 and J4, nor in migratory females; these stages apparently do not feed. Observations suggest that reserves stored by J2 sustain three ecdyses and the migratory female's search for a feeding site and induction of a syncytium. Feeding activity is resumed in young and mature sedentary females, in which the DG is highly active and enlarged. The SvG are metabolically active, but with little synthesis of secretory granules, suggesting that in sedentary females the SvG may have physiological roles other than digestion.
esophageal glands; feeding biology; gland activity; Nacobbus aberrans; nematode; Pratylenchidae; ultrastructure
Cactodera salina n. sp. (Heteroderinae) is described from roots of the estuary plant Salicornia bigelovii (Chenopodiaceae), in Puerto Pefiasco, Sonora, Mexico, at the northern tip of the Sea of Cortez. The halophyte host is grown experimentally for oilseed in plots flooded daily with seawater. Infected plants appear to be adversely affected by C. salina relative to plants in noninfested plots. Cactodera salina extends the morphological limits of the genus. Females and cysts have a very small or absent terminal cone and deep cuticular folds in a zigzag pattern more typical of Heterodera and Globodera than of Cactodera spp. Many Cactodera spp. have a tuberculate egg surface, whereas C. salina shares the character of a smooth egg with C. amaranthi, C. weissi, and C. acnidae. Only C. milleri and C. acnidae have larger cysts than C. salina. Face patterns of males and second-stage juveniles, as viewed with scanning electron microscopy, reveal the full complement of six lip sectors as in other Cactodera spp. Circumfenestrae of C. salina are typical for the genus.
Cactodera salina; cyst nematodes; halophyte; Heteroderinae; nematode; new species; Salicornia bigelovii; scanning electron microscopy; Sea of Cortez; taxonomy
Biochemical and molecular analyses of genetic variation were evaluated to address the taxonomic status of Nacobbus aberrans. Isolates from Mexico, Peru, and Argentina, cultured on tomato in the greenhouse, were analyzed with respect to isozyme and DNA marker variation. Although acid phosphatase and malate dehydrogenase revealed distinct profiles for each isolate, non-specific esterases revealed possible affinities between the Peruvian isolates and between the isolates from Mexico and Peru. Two of l 0 RAPD primers revealed affinities suggested by esterase profiles. RFLP analysis of the rDNA repeating unit with six restriction enzymes revealed identical cleavage patterns between the Peru isolates and a distinct profile shared by isolates from Mexico and Argentina. Nucleotide sequence analysis of the 5.8S rRNA coding region revealed differences among the four isolates at eight of 157 positions; sequences of the Peruvian isolates differed from each other at only one position, whereas the Mexican and Argentine isolates were identical and could be distinguished from the Peruvian isolates. A distance matrix from unweighted pairwise comparisons of the 5.8S rDNA revealed apparent elevated intraspecific divergence in N. aberrans comparable to intergeneric divergence between Heterodera and Globodera. Analysis of additional N. aberrans isolates from throughout the distribution range should help determine the full extent of intraspecific genetic variation that underlies the phenotypic and morphologic diversity of the genus.
esterase; false root-knot nematode; molecular biology; Nacobbus aberrans; nematode; RAPD; rDNA; RFLP; 5.8S rRNA; taxonomy
Meloidogyne sp. from five pecan (Carya illinoensis) orchards in Texas were distinctive in host range and iszoyme profiles from common species of Meloidogyne but were morphologically congruent with Meloidogyne partityla Kleynhans, a species previously known only in South Africa. In addition to pecan, species of walnut (Juglans hindsii and J. regia) and hickory (C. ovata) also were hosts. No reproduction was observed on 15 other plant species from nine families, including several common hosts of other Meloidogyne spp. Three esterase phenotypes and two malate dehydrogenase phenotypes of M. partityla were identified by polyacrylamide gel electrophoresis. Each of these isozyme phenotypes was distinct from those of the more common species M. arenaria, M. hapla, M. incognita, and M. javanica.
Carya illinoensis; C. ovata; detection; esterase; hickory; isozyme phenotype; Juglans hindsii; J. regia; malate dehydrogenase; Meloidogyne partityla; pecan; root-knot nematode; walnut
Fine structure of developing sperm of the monospecific genus, Ekphymatodera, was compared with other Heteroderinae as part of a study to recognize diversity and phylogenetically informative characters within the subfamily. Sperm of Ekphymatodera originate from germ cells connected to a central rachis, a character which is shared with Globodera, but not with other Heteoderinae. In Ekphymatodera, and cyst-forming genera, a layer of cortical microtubules lies just beneath the surface of the plasma membrane. Sperm of Ekphymatodera are unique among Heteroderinae examined by the presence of spiral surface elevations on the filopodia, a character that may prove to be a synapomorphy for Sarisoderini. Fibrous bodies are abundant in spermatids; however, they do not persist in sperm of Ekphymatodera as they do in Meloidodera and Verutus. The male gonad of Ekphymatodera is lined by epithelial cells, which are greatly enlarged near the ejaculatory canal. These enlarged cells contain vesicles with concentric lamellar inclusions, not observed in other genera of the subfamily. Sperm of Heteroderinae are rich in diversity, and examination of additional representative species may indicate new phylogenetically informative characters.
Ekphymatodera thomasoni; filopodia; fine structure; Heteroderinae; lamellar inclusion; male gonad; nematode; ontogeny; phylogeny; pseudopodia; sperm; systematics; ultrastructure
Bursaphelenchus abruptus n. sp., an associate of the digger bee, Anthophora abrupta (Hymenoptera: Anthophoridae), is described and illustrated. Bursaphelenchus abruptus n. sp. can be differentiated from other species of Bursaphelenchus by the absence of head annules, stylet length, length of the postuterine sac, shape of female tail, spicule morphology, and male caudal papillae arrangement. Two plant-pathogenic fungi, Monilinia fructicola and Botrytis cinerea, and a Monilia sp. isolated from an adult bee from Prince Georges County, Maryland, were good hosts for B. abruptus n. sp. Dauer juveniles (JIII) of B. abruptus n. sp. were isolated from the reproductive tracts of A. abrupta from Montgomery County, Alabama, for measurements and comparison with J2 -JIII inter-molts from a 4-week-old monoxenic culture on Monilia sp. Gonad lengths in dauer juveniles isolated from A. abrupta were highly variable (49 ± 23 μm SD; range 21-93 μm; n = 29) compared with J2-JIII intermolts from culture (28 ± 7 μm SD; range = 16-42 μm; n = 16), suggesting that postembryonic gonad development may continue while dauers are in the bee host. Adult males and females of B. abruptus n. sp. were examined with scanning electron microscopy (SEM) for ultrastructural comparisons with other members of the genus Bursaphelenchus.
Anthophora abrupta; Aphelenchoididae; bee; Bursaphelenchus abruptus n. sp.; B. fraudulentus; B. kolymensis; B. mucronatus; B. xylophilus; Dufour's gland; morphology; mycophagy; nematode; scanning electron microscopy; taxonomy
Fine structure of the posterior cone of monoxenically cultured Heterodera schachtii is examined. The cone is not evident at the end of the fourth molt, but as the female matures the cone elongates, vulval lips enlarge, and cuticular patterns on the lips are modified. Body wall cuticle (BW) of the cone includes layers A and B, but C is modified or replaced by a network of fibers which correspond to the semifenestrae. Vaginal lining is continuous with the BW and terminates at the cuticular underbridge near the uterus. Vaginal musculature includes 48 dilatores vaginae (DV) as well as a sphincter vaginae (SV). The DV include a contractile and noncontractile region with abundant actin and glycogen. A distinct anal depressor muscle is present. In the cyst, only bullae, the underbridge, vagina lining, and traces of the SV muscle persist. Detailed morphology of the cone of H. schachtii provides insight into characters which, when compared with other heteroderines, will be useful in phylogenetic analysis of Heteroderinae.
cone; cyst; dilatores vaginae; fenestrae; fluorescent phalloidin; Heterodera schachtii; monoxenic culture; phylogeny; scanning electron microscopy (SEM); sphincter vaginae; transmission electron microscopy (TEM); underbridge; vagina
Phylogenetic characters for Heteroderinae Luc. et al., 1988 are evaluated in Meloidodera which is believed to have primarily ancestral characters. Phasmid ultrastructure is observed in second-stage juveniles (J2), third-stage juvenile males, fourth-stage juvenile males, and fifth-stage males of Meloidodera floridensis and M. charis. Phasmid secretion occurs inside the egg before the J1-J2 molt. Before J2 hatch, concentric lamellar membranes occur within the sheath and socket cells. Some membranes become lamellae of the sheath cell plasma membrane; others become multilamellar bodies. During early molting, plasma membrane lamellae disappear and a distal dendrite segment appears in a rudimentary canal. After the molt, the distal dendrite is not present within the canal. The phylogenetic utility of phasmid features is discussed. In both species the ampulla shape and size between molts are stable features in juveniles and males. The posthatch J2 sheath cell receptor cavity may vary in a species specific manner, but comparative morphology requires precise timing after hatch.
end apparatus; Heteroderinae; male development; Meloidodera floridensis; Meloidodera charis; molting; multilamellar body; neuromorphology; phasmid; phylogeny; secretion; ultrastructure
Fine structure of the body wall cuticle of Heterodera schachtii is compared with respect to age and body region of the female. The cuticle is more complex than previously reported. In newly molted females only layers A, B, and C are present, but 4 weeks after the final molt a thin D layer is present between the midbody and base of the cone. This D layer is absent in the cone of H. schachtii, regardless of age. As females age, an additional layer E is produced and includes zones E₁ and E₂. Zone El apparently is unique to H. schachtii, whereas E₂ is likely to be homologous with a similar layer in Atalodera. In the cone of old females (ca. 8 weeks after the final molt) of H. schachtii, the two zones become irregular in shape and comprise bullae. The presence of a thin D layer in Heterodera strengthens the previous hypothesis of a single ancestor of cyst nematodes.
cone; cuticle; cyst; Heterodera; Heteroderinae; phylogeny; transmission electron microscopy (TEM)
Absence of the phasmid was demonstrated with the transmission electron microscope in immature third-stage (M3) and fourth-stage (M4) males and mature fifth-stage males (M5) of Heterodera schachtii, M3 and M4 of Verutus volvingentis, and M5 of Cactodera eremica. This absence was supported by the lack of phasmid staining with Coomassie blue and cobalt sulfide. All phasmid structures, except the canal and ampulla, were absent in the postpenetration second-stage juvenile (J2) of H. schachtii. The prepenetration V. volvingentis J2 differs from H. schachtii by having only a canal remnant and no ampulla. This and parsimonious evidence suggest that these two types of phasmids probably evolved in parallel, although ampulla and receptor cavity shape are similar. Absence of the male phasmid throughout development might be associated with an amphimictic mode of reproduction. Phasmid function is discussed, and female pheromone reception ruled out. Variations in ampulla shape are evaluated as phylogenetic character states within the Heteroderinae and putative phylogenetic outgroup Hoplolaimidae.
anaphimixis; ampulla; cell death; Cactodera eremica; Heterodera schachtii; Heteroderinae; parallel evolution; parthenogenesis; phasmid; phylogeny; ultrastructure; Verutus volvingentis
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.
Atalodera trilineata n. sp.; A. festucae n. sp.; Ekphymatodera n. gen.; E. thomasoni n. sp.; Heteroderidae; new genus; new species; scanning electron microscopy; taxonomy; Verutus californicus n. sp.
SEM examination of second-stage juveniles (J2) and adults of Atalodera ucri, A. lonicerae (syn. Sherodera lonicerae), Thecavermiculatus sp. (undescribed new species), T. andinus, and T. crassicrustatus revealed new characters. A primitive en face pattern with six separate lips occurs in J2 of Thecavermiculatus spp. examined and in about half the polymorphic A. lonicerae. A derived en face pattern with fused adjacent submedial lips occurs in the other half of A. lonicerae and all A. ucri. Posteriorly, the J2 head of all species is annulated. The primitive en face pattern also occurs in males of A. lonicerae and Thecavermiculatus spp., and posteriorly the head of these species consists of plates. Fewer plates occur rarely in males of A. ucri. Males of A. ucri have a derived en face pattern where lips are fused and the head is annulated. Fusion of lips occurs rarely in males of A. lonicerae. Females of all species have similar derived en face patterns. En face patterns of J2 and males o f Atalodera and Thecavermiculatus may aid in species identification and to elucidate intergeneric relationships, but en face characters shared by the two genera are primitive and are not useful for demonstrating monophyly. Perineal region of females indicates the closeness of the vulval-anal distance, as a derived character, which is shared by Atalodera and most Thecavermiculatus spp. suggesting possible monophyly. T. andinus, while having a similar en face pattern to J2 of other Thecavermiculatus species, lacks the derived character of the perineal region. Phasmid openings were not observed in adults of any of the species examined.
en face pattern; perineal region; phasmid; phylogeny; SEM; systematics; vulva
Scanning electron microscopy (SEM) of second-stage juveniles (J2), males, and females of Meloidodera floridensis, M. charis, M. belli, and Verutus volvingentis reveals detailed characteristics of the head region, lateral field, phasmid, body striae, vulva, and perineal region. In M. charis and M. belli the en face pattern conforms to a basic pattern in which the labial disc is surrounded by six lips (sectors) of the first head annulation. In J2 the head has additional annulations, whereas in males annulation is replaced by longitudinal blocks. Conversely, J2 and males of M. floridensis and V. volvingentis each have a unique derived face pattern with fusion of various lip components and with head annulation. All six lips of females of M. charis and M. belli are fused, whereas females of M. floridensis and V. volvingentis have distinct lateral lips. Lateral fields vary among species, with only slight differences at the anterior and posterior ends of the lateral lines and in the spatial relation of the lines to phasmid openings. Phasmid openings are present in adults of Meloidodera spp., but were not observed in adults of V. volvingentis; in this respect, the female perineal pattern of Verutus is different from Meloidodera spp, The very large vulva (± 48 μm long) of V. volvingentis is in sharp contrast to the minute vulva (± 6 μm long) in a population of M. charis from San Bernardino. Morphological characters revealed by SEM will be most informative when investigated throughout Heteroderidae and incorporated with additional characters for a phylogenetic analysis of the family.
en face patterns; perineal pattern; phasmids; phylogeny; SEM; systematics; vulva
Body wall cuticle of adult females of eight genera within the Heteroderidae was examined by transmission electron microscopy for comparison with previously studied species within the family. Cuticle structure was used to test some current hypotheses of phylogeny of Heteroderidae and to evaluate intrageneric variability in cuticle layering. Verutus, Rhizonema, and Meloidodera possess striated cuticle surfaces and have the simplest layering, suggesting that striations have not necessarily arisen repeatedly in Heteroderidae through convergent or parallel evolution. Atalodera and Thecavermiculatus possess similar cuticles with derived characteristics, strengthening the hypothesis that the two genera are sister groups. Similarly, the cuticle of Cactodera resembles the specialized cuticle of Globodera and Punctodera in having a basal layer (D) and a surface layer infused with electron-dense substance. Heterodera betulae has a unique cuticle in which the thickest layer (C) is infiltrated with an electron-dense matrix. Little intrageneric difference was found between cuticles of two species of Meloidodera or between two species of Atalodera. However, Atalodera ucri has a basal layer (E) not found in other Heteroderidae. The most striking intrageneric variation in cuticle structure was observed between the thin three-layered cuticle of Sarisodera africana and the much thicker four-layered cuticle of Sarisodera hydrophila; results do not support monophyly of Sarisodera.
Atalodera ucri; body wall cuticle; Cactodera sp.; comparative morphology; fine structure; Heterodera betulae; Heteroderidae; Meloidodera floridensis; phylogeny; Rhizonema sequoiae; Sarisodera africana; systematics; Thecavetmiculatus gracililancea; Verutus volvingentis
Systematic contributions to Heteroderidae include description of Cactodera eremica n. sp., an emended diagnosis of Sarisodera Wouts and Sher, 1971, and proposal of a new genus and new combination, Afenestrata africana (synonym Sarisodera africana Luc et al., 1973). Cactodera eremica, from the roots of shadscale in Utah, most closely resembles Cactodera thornei (Golden and Raski, 1977) but differs by the presence of a finely striated cuticle, a fine surface pattern on eggs, a shorter female stylet, distance of the DGO from the stylet, vulval slit, and smaller diameter of circumfenestra, as well as a shorter tail in second-stage juveniles. The response of the host to C. eremica is similar to other Heterodera sensu lato including a large syncytium with wall ingrowths. The diagnosis of Sarisodera is emended to exclude cysts, which do not form in the type species, S. hydrophila. Afenestrata africana differs from S. hydrophila by the formation of cysts, the dorsal position of the anus in females, the shorter stylet, and a pore-like phasmid opening in second-stage juveniles. In addition, the lip pattern of males and juveniles is characterized by a greater degree of fusion of lip parts, the host response is a syncytium (versus a single uninucleate giant cell in S. hydrophila), and the cuticle is thinner and lacks a D layer. Unlike Heterodera, the cyst of Afenestrata lacks fenestrae.
comparative morphology; cyst nematodes; giant cell; Heterodera; histopathology; host response; scanning electron microscopy; taxonomy
junior primary homonym; Tylenchorhynchus
Host responses to Meloidodera floridensis Chitwood et al., 1956, M. charis Hooper, 1960, and M. belli Wouts, 1973 were examined on loblolly pine, peony, and sage, respectively, with light, scanning, and transmission electron microscopy. In each case the nematodes induce a single uninucleate giant cell. The giant cell is initiated in the pericycle and expands as it matures. The mature giant cell induced by M. floridensis is surrounded by vascular parenchyma, whereas that caused by M. charts and M. belli coutacts xylem and phloem. The cell wall of giant cells induced by all three Meloidodera spp. is generally thicker than that of surrounding cells, with the thickest part adjacent to the lip region of the nematode. The thinner portion of the wall includes numerous pit fields with plasmodesmata, but wall ingrowths were not detected in a thorough examination of the entire wall. The nucleus of a giant cell induced by M. goridensis is highly irregular in shape with deep invaginations, whereas those caused by M. charis and M. belli include a cluster of apparently interconnected nuclear units. Organelles, including mitochondria, endoplasmic reticulum, and plastids of giant cells caused by Meloidodera, are typical of those reported in host responses of other Heteroderidae. The formation of a single uninucleate giant cell by Meloidodera, Cryphodera, Hylonerna, and Sarisodera, but a syncytium by Atalodera and Heterodera sensu lato, might be considered in conjunction with additional characters to determine the most parsimonious pattern of phylogeny of Heteroderidae.
callose; giant cell; Heteroderoidae; histopathology; plasmodesmata; wall ingrowths
The body wall cuticle of adult females of Meloidodera charis, Atolodera lonicerae, and Sarisodera hydrophila is examined by transmission electron and light microscopy for comparison with Heterodera schachtii and previous observations of additional species of Heterodera, Globodera, and Punctodera. The cuticle of M. charis is least complex, consisting of layers A, B, C (with A outermost), and varies in overall thickness from 3 to 8 μm. As in other species, the cuticle is thickest in mature specimens. The cuticle of A. lonicerae is 6-9 μm thick; unlike M. charis it has an innermost layer, D, in addition to A, B, and C. The cuticle of S. hydrophila varies from 14 to 30 μm thick and includes a D layer similar to A. lonicerae; layer C is subdivided into additional zones relative to other heteroderids, and the external portion of the cuticle is infused with an electron-dense material. The presence of a D layer in A. lonicerae and S. hydrophila is a character state which is shared with Globodera spp. and Punctodera sp. The electron-dense material in the outer layers of S. hydrophila also occurs in Globodera spp. and Punctodera sp. On the other hand, H. schachtii resembles other Heterodera spp. as well as M. charis by the absence of a D layer and lack of electron-dense material in the outer layers. The pattern of occurrence of shared character states, including those of the cuticle, may be useful for phylogenetic analysis of Heteroderidae.
cyst; electron microscopy; Globodera; Heterodera; Punctodera; Heteroderoidea; Tylenchida; phylogeny