Search

Forty-four specimens of the millipede Narceus gordanus Chamberlin, 1943 (Spirobolida: Spirobolidae) were collected from Alachua, Citrus, Hernando, and Marion counties in peninsular Florida. Morphometric data were recorded for each. Nematodes were dissected from the intestine of each individual and sorted into morphotaxa. Heth pivari n. sp. (Oxyuridomorpha: Ransomnematoidea: Hethidae) was found in 33 (75%) of dissected N. gordanus and examined with brightfield, differential interference contrast, phase contrast and scanning electron microscopies. LSU rDNA sequences of representative males and females of H. pivari n. sp. were analyzed and compared to sequences of nematodes in the infraorder Rhigonematomorpha. Heth mauriesi, an introduced species, also was sequenced. Females of H. pivari n. sp. differ from those of other Heth spp. in having smooth, button-like somatic and cervical papillae and shallow, shield-like cervical collars. Males have slit-like or narrowed, rather than circular, stomal openings. Heth pivari n. sp. is the first species of this genus found in an indigenous millipede north of Mexico. Keys based on female cervical ornamentation are provided to differentiate the 52 known Heth spp.

Heth pivari n. sp. (Tables 2–4; Figs. 2–7) Type locality and habitat. Florida: Marion County, Silver Springs, Ocala National Forest (29.257726 N, – 81.778702 W), xeric scrub located near Mount Dora Sand Ridge, in intestine of Narceus gordanus. Other specimens collected: Ridge Manor, Hernando County (28.509794 N, – 82.186692 W); Citrus Springs, Citrus County (29.9786111 N, – 82.4327777 W); and Gainesville, Alachua County (28.707009 N, – 89.395251 W), all in N. gordanus. Type designation and deposition. Holotype female (T- 708t) and male paratype (T-6937p), deposited in the USDA Nematode Collection, Beltsville, Maryland. Additional specimens deposited in the nematode collection in the Entomology and Plant Pathology Department, University of Tennessee, Knoxville, Tennessee (University of Tennessee Control Numbers B 1–B24). Etymology. We are pleased to dedicate this new species to Dr. Robert J. Pivar, outstanding former graduate student in the Entomology and Plant Pathology Department, University of Tennessee, Knoxville. Females. Measurements and ratios given in Table 2. Body cylindrical, white in life, robust, anterior head region broad; maximum diameter near mid-body, then tapering posteriorly and terminating in filiform tail (Figs. 2A, 7A). Differentiated lateral field absent. Annules 1–1.5 µm wide posterior to cervical collar, each annule from neck to anal region with numerous minute longitudinal striae (Fig. 3D); neck region finely annulated but with weak or no transverse striations (Figs. 3 A–C). Buccal cavity tubular, expanded anteriorly, much detail obscured by other head structures; anterior and posterior parts divided by circular ring of short, longitudinal striae; posterior part with extremely fine transverse striae about 0.5 µm apart (Figs. 2B, C). Amphid apertures circular, 2–4 µm in diameter, located on convex lateral aspect of each pseudolabium (Figs. 3 A–C). Pseudolabia rectangular in dorso-lateral view (Fig. 3A), rounded on outer margin in dorsal view (Figs. 3 A-C), ornamented with combs and spines; small pectinate combs with 2–3 µm-long bristles on interior lateral, median and cleft margins, and larger combs with 5–7 µm-long bristles continuing along dorsoventral aspect of each pseudolabium; dorsal and ventral bristles rounded distally; length of bristles on each margin approximately equal (Fig. 3C). Anterior stomal region often filled with amorphous material (Fig. 3B). Neck region with four folds anterior to cervical collar, with second, third and fourth folds each containing one smooth, knob-like cervical sensory papilla per quadrant (Figs. 3B, C); papillae about 1–2 µm in diameter. Cervical collar with about 72 spines, 5–6 µm long; collar interrupted laterally on each side by shield, edge spines often bi- or trifurcate (Figs. 3A, D); shield wider than long (Figs. 2D, 3A, 6A). Two lateral pairs of large, acute spines in tandem, 10–13 µm long; anterior pair sometimes connected by a shallow ridge, posterior pair less frequently so (Figs. 3A, B). Annulation between each pair of spines irregular and areolated. Knob-like somatic papillae scattered along length of body, denser anterior to vulva (Figs. 3D, 6D). Neck region dorsally and ventrally with retractor muscles attached to body wall at each major fold, integrated with strong longitudinal muscle associated with large nucleus (Figs. 3B, C). Two-part esophagus consisting of procorpus and basal bulb with grinding valve, surrounded anteriorly by six prominent, uninucleate, amber-colored glands (Fig. 2A) identical in number and arrangement to those of male (Figs. 4A, B, 6B) and one or more small colorless glands (Fig. 2B). Secretory-excretory (SE) pore minute, inconspicuous, 167–234 µm from anterior head (Figs. 2A, 6C); SE complex fused with nerve ring and anchored dorsally to body wall, identical to that of male (Fig. 4B). Four coelomocytes: one in esophageal region, two next to anterior region of intestine or near anterior regions of ovaries, one near spermathecae (Fig. 2A). Lateral nerve chord clearly visible, identical to that of male (Fig. 7H). Two gonads, pro-didelphic, reflexed; spermathecae present, filled with large, fusiform or comet-shaped motile sperm (Figs.2A, E, 6E). Muscular, spherical sphincter at junction of gonads with oviduct. Uterus prominent, distinctly muscled (Figs. 2E, 7C, E). Eggs large, few to moderate in number, eggshells thin, without ornamentation (Figs. 7 A–E). Vulva near anus. Phasmid apertures minute, pores located just posterior to anus. Tail filiform, attenuated to a fine point. Males. Measurements given in Table 3. Body robust, smaller than females, white, head and neck without ornamentation (Figs. 4A, 5A, B, C); lateral field absent; cuticle finely annulated, each annule 1–1.5 µm wide, with scattered, smooth papillae (Figs. 5B, E). Stomatal opening longer than wide to slit-like (Figs. 4B, C, 5 A–C). Stoma tubular, with three feather-like appendages (Fig. 5B); stomatal structure partially obscured by tapering, partially striated esophageal collar. Lips finely striated, slightly elevated with four inconspicuous cephalic papillae (Figs. 5B, C). Amphid apertures inconspicuous, circular, on inner slope of raised lip region, approximately 3 µm in diameter (Fig. 5C). Head region with pairs of subdorsal and subventral muscles attached to stoma and extending obliquely to body wall (Figs. 4B, C). Six prominent amber-brown, uninucleate gland-like somatic extensions of posterior arcade syncytium surrounding anterior part of esophagus and extending to nerve ring (Figs. 4B, 6B); at least two much smaller, colorless extensions of the anterior arcade syncytium located near base of stoma (Fig. 4B). Esophagus approximately onethird of body length, procorpus muscled; broadly pyriform basal bulb containing grinding valve. Intestine dilated at anterior end, then attenuating to a uniform diameter and in ventral position, crossing testis near ventral sucker and becoming dorsal, then narrowing at cloaca (Figs. 4A, D). Secretory-excretory (SE) complex and nerve ring forming a single arching structure (Fig. 4B); SE pore inconspicuous, connected to sinuous tube in the SE complex; nerve ring thin, with many associated nuclei, fused with connective tissue extending to dorsal cuticular wall (Fig. 4B). Four coelomocytes, one each in esophageal region, in anterior area of intestine, near flexure of testis and near junction of seminal vesicle and vas deferens. Lateral nerve chords clearly visible (Fig. 7H). Reproductive system monorchic, with short flexure in germinal zone. Sperm comet to fusiform-shaped, about 88 µm long × 15 µm at widest point (Fig. 6E). Ventral sucker present, 202–330 µm anterior to cloacal opening, flanked by first pair of large pre-cloacal papillae (Figs. 4D, 5D). Region between sucker and cloaca with scattered somatic papillae of 1–2 µm diameter. Spicules equal, fused for distal three-fourths; each spicule with distinct capitulum, apical fused portion arcuate ventrad, tapering to a point; gubernaculum with pointed proximal end, distal end broad and bilobed (Figs. 4D, E). Cloacal lips smooth. Seven pairs of genital papillae, each with several conspicuous internal striae or minute tubes; one median ventral papilla. Caudal papilla pattern as follows: first pair slightly posterior to ventral sucker; second pair about 30 µm anterior to cloaca, distant from third pair; pairs 3 and 4 slightly anterior to cloaca, pair 3 ventral, pair 4 lateral; single medial papilla (5) on anterior cloacal lip; posterior to cloaca, two ventral pairs (6, 7) and one subdorsal pair (8) (Figs. 4D, 5 D–F). Differential diagnosis. Females of Heth pivari n. sp. are characterized by smooth, knob-like cervical and somatic papillae, shallow cuticular shields, continuous cuticular collars with approximately 72 subequal spines, and two pairs of paired anterior and posterior lateral spines. Males lack ornate cuticular ornamentation, have a narrowed stomal opening and smooth somatic papillae, but otherwise are similar to other described males. The morphology of females of H. pivari n. sp. is most similar to those Heth spp. having a continuous collar of cervical spines with a spiny lateral shield. Heth pivari n. sp. is distinct from H. mauriesi Adamson, 1982, its closest geographic neighbor, in female length (2,190 –4,483 µm vs. 1,575 –2,000 µm in H. mauriesi), smooth cervical and somatic papillae (multi-cusped in H. mauriesi), and a trapezoidal shield interrupting the cervical collar (shield replaced by two stout spines in line with the smaller collar spines in H. mauriesi) (Figs. 1A, B). Among the other subtropical and tropical North American species, H. tuxtlensis (Fig. 12B) is most similar to H. pivari n. sp. in having a continuous cervical collar, the presence of lappets that usually do not overlap the anterior lateral spines, absence of lateral alae, and a finely striated body. Heth pivari n. sp. and H. tuxtlensis are easily differentiated by the number of cervical spines (ca. 72 for H. pivari n. sp., ca. 100 for H. tuxtlensis), a shallow cervical shield in H. pivari n. sp., (more pronounced in H. tuxtlensis), more bifurcated lappet spines in H. tuxtlensis as compared to fewer cervical spine bifurcations in H. pivari n. sp. and smaller anterior and posterior lateral spines (10–13 µm long) compared to the larger and more robust spines of H. tuxtlensis (anterior spine length 58–67 µm, posterior spine length 82–94 µm posterior spines). Females of H. pivari n. sp. resemble those of H. taybaci (Vietnam) in that the shields have similar shapes. The posteriormost spines of the cuticular collar nearly reach the base of the anterior lateral spines, and both have smooth somatic papillae. Heth pivari n. sp. has fewer collar spines (approximately 72) compared to about 88 around the circumference of H. taybaci. Heth pivari n. sp. also differs in that the anterior and posterior lateral spine pairs are smaller than those of H. taybaci. Males of H. pivari n. sp. have smaller spicule arcs (mean 109 µm) than H. taybaci (136 µm) but have longer bodies (Heth pivari n. sp. males 1,897 –2,609 µm, H. taybaci 1,520 µm). Two additional Heth species with a continuous shield-like collar occur in the western hemisphere: H. insularis (Brazil) and H. orthopori (Paraguay). In both of those species the lateral spine pairs are broadly joined at their bases. All other Heth species in this group are located in Australia, New Zealand, and Asia. Heth pivari n. sp. has short lappets whose posteriormost spines do not reach the anterior lateral spines, whereas in H. costata, H. impalutiensis, H. sutherlandi, H. taynguyeni, and H. vietnamensis the spines of the much longer lappets overlap the anterior lateral spines. Heth baudini and H. dimorphum have shields in which length and width are about equal, whereas in H. pivari n. sp. the shallow shield width is about twice its length. The type species, H. juli, has a “W” shape as opposed to a trapezoidal shape in H. pivari n. sp. Finally, H. xaniophora has prominent lateral alae and robust anterior and posterior lateral spines, whereas H. pivari n. sp. lacks lateral alae and has small anterior and posterior lateral spines. Molecular analysis. With the addition of H. pivari n. sp. and H. mauriesi, nine Heth species were included in a 28S rDNA tree (Fig. 8). Heth pivari n. sp. and two Mexican species, H. gordae and H. xarochae, grouped together with high support. The other six, including H. mauriesi, had no support except for H. initiaensis - H. konoplevi. The molecular relationship of H. pivari n. sp. to H. gordae and H. xarochae is supported by geographical location but not by morphological similarity., Published as part of Phillips, Gary, Moulton, John K. & Bernard, Ernest C., 2020, Heth pivari n. sp. (Nematoda: Ransomnematoidea: Hethidae) from the indigenous North American millipede Narceus gordanus (Spirobolida: Spirobolidae), with keys for worldwide Heth spp., pp. 486-514 in Zootaxa 4861 (4) on pages 492-498, DOI: 10.11646/zootaxa.4861.4.2, http://zenodo.org/record/4416659, {"references":["Spiridonov, S. E. & Yushin, V. V. (2000) Ultrastructure of the stoma in Heth mauriesi Adamson, 1982 (Rhigonematida: Hethidae). Nematology, 2, 417 - 424. https: // doi. org / 10.1163 / 156854100509277","Adamson, M. L. (1982) Two new species of Heth Cobb, 1898 (Nematoda, Rhigonematidae), from South American diplopods. Bulletin de Museum National de Histoire Naturelle 4 e Serie Section, A 4, 405 - 418."]}

Key to group 1 Heth spp. 1. With one continuous cervical collar of spines around circumference of neck, usually with interruption by lateral cervical spines.............................................................................................. 2 - With two continuous cervical collars around circumference of neck; rows of eight similar-sized spines extending from cervical collar to level of esophageal isthmus (Fig. 9E); lateral alae originating from posteriormost cervical spines and ending at base of tail; female length 1,326 –1,440 µm.................................................................. H. coyi 2. Lateral spines absent; cervical collar spines continuous, not interrupted laterally; numerous multi-cusped sensory papillae anterior and posterior to cervical collar (Fig. 9A); length of female 1,070 –1,210 µm.......................... H. albertoi - Lateral spines present; cervical collar interrupted or deeply notched laterally, often by prominent cervical spines.......... 3 3. One or two pairs of lateral spines present, posterior lateral spines sometimes single................................. 4 - Numerous lateral spines (>8 pairs) extending half the length of the esophagus; lateral alae wing-like.................. 15 4. Single posterior lateral spine............................................................................ 5 - Posterior lateral spines paired or absent.................................................................... 7 5. Cervical collar interrupted laterally (Fig. 10G); length of female 1,550 –1,920 µm...................... H. zeuglocantha - Cervical collar notched or slightly produced laterally as two small, pointed lobes................................... 6 6. Cervical collar notched (Fig. 9I); female length 1,730 -2,240 µm....................................... H. konoplevi - Cervical collar slightly prolonged as two small, pointed lobes (Fig. 11D); females 1,420 –2,500 μm long....... H. hamatus 7. Lateral spine pairs broad, outer edges coarsely toothed....................................................... 8 - Lateral spine pairs slender, outer edges smooth............................................................. 9 8. Cervical collar with about 92 spines; two broad, strongly serrated pairs of lateral spines, their length and width about equal; length of female 2,600 –3,200 µm (Fig. 9F)......................................................... H. gordae - Cervical collar with about 60 spines; anterior and posterior lateral spines much wider than long, posterior spines nearly encircling the body; female length 1,400 –1,700 µm (Fig. 10F)............................................. H. xarochae 9. With continuous anterior and posterior series of discontinuous spines around the circumference of the esophageal region (Fig. 9J); length of female 1,731 –2,482 µm........................................................... H. maicuru - Post-collar spines confined to lateral region............................................................... 10 10. Cervical collar densely serrated and expanded away from body; posterior spines apparently absent; length of female 1,820 – 2,070 µm (Fig. 9G)....................................................................... H. hexaspinosum - Cervical collar not expanded away from body; posterior lateral spines present.................................... 11 11. Large multi-cusped papillae (2–4 per side) only present anterior to cervical collar................................. 12 - Papillae present both anteriorly and posteriorly to cervical collar; with transverse rows of small spines in cervical folds... 13 12. Cervical, anterior and posterior lateral spines narrow, about equal in size (Fig. 10B); length of holotype 2,800 µm................................................................................................ H. perarmatum - Cervical, anterior and posterior lateral spines progressively larger, broader and overlapping one another (Fig. 9B); lateral winglike alae extending from posterior region of esophagus to anus; female length 1,753 –1,973 µm.......... H. amazonensis 13. Collar interrupted by long, slender cervical spines nearly reaching base of anterior lateral spines; anterior and posterior lateral spines long and slender (Fig. 10E); female length 1,610 –1,990 µm.................................... H. spinosum - Cervical collar interrupted by larger lateral spines not reaching base of anterior lateral spines........................ 14 14. Anterior and posterior spine pairs not sharing common bases (Figs. 1, 10A); female length 1,750 –1,830 µm.... H. mauriesi - Anterior and posterior spine pairs with common bases....................................................... 15 15. Anterior lateral spines not reaching base of posterior spines (Fig. 9D); female length 2,960 –3,090 µm........... H. clunyi - Anterior pair of lateral spines overlapping base of posterior pair; (Fig. 9C); female length 1,880 –2,080 µm H. bifidispiculum 16. Prominent spines extending from posterior cervical collar to beginning of lateral alae (Fig. 10C, D); lateral alae extending from posterior esophagus to tail; length of female 1,155 –1,165 µm........................................ H. pinnatum - Somatic spines extending from cephalic extremity to isthmus with first three spines small and remaining spines long and curved (Fig. 9H); female length 1,740 –2,000 µm...................................................... H. hispaniolae, Published as part of Phillips, Gary, Moulton, John K. & Bernard, Ernest C., 2020, Heth pivari n. sp. (Nematoda: Ransomnematoidea: Hethidae) from the indigenous North American millipede Narceus gordanus (Spirobolida: Spirobolidae), with keys for worldwide Heth spp., pp. 486-514 in Zootaxa 4861 (4) on pages 506-507, DOI: 10.11646/zootaxa.4861.4.2, http://zenodo.org/record/4416659