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Diphtherophora species phylogeny In total, 26 DNA sequences, specifically covering the D2���D3 domains of the 28S rRNA gene, were used in our molecular phylogenetic analyses: 18 Diphtherophora, two Tylolaimophorus, one representative of Prismatolaimidae, six representatives of Trichodoridae, two representatives of Tripylidae, and three representatives of Trischistomatidae. Our final alignment was 785 bp long and contained 326 conserved sites, 455 variable sites, 415 parsimonyinformative sites, and 40 singletons. Phylogenetic analyses based on both inference methods (i.e., ML and BI) were highly congruent with respect to the monophyly of the genus Diphtherophora, the relationships among Diphtherophora species and outgroups, and the phylogenetic placement of D. eldarica n. sp. (Fig. 4). Phylogenetic trees showed the existence of six strongly supported clades (A���F) within the Diphtherophora group: clade A is represented by D. cf. caudata from Arasbaran (Northwestern Iran) and D. caudata from Maragheh (Northwestern Iran), the latter species recovered in this study; clade B included sequences representing D. cf. obesa and D. tenera from different regions of Northern and Southern Iran, the latter species, however, being paraphyletic; clade C was comprised of D. perplexans from different sites in Northwestern (Takab and Urmia) and Southern Iran (Shiraz), the latter population recovered in this study, and a specimen characterized only as Diphtherophora sp. from CA, USA; clade D was solely represented by the new species D. eldarica n. sp.; clade E harbored uncharacterized Diphtherophora species from different sites in Northern and Northwestern Iran including Arasbaran and Ali Abad; clade F is solely represented by sequences of D. geraerti from Shiraz (Southern Iran). Our phylogenetic analyses strongly supported (ML = 97, BI = 1.0) the new species D. eldarica n. sp. as a sister taxon of clade D, that is, D. perplexans + Diphtherophora sp. (Fig.4). Genetic divergence (p-distance %) within Diphtherophora clades was relatively low, ranging from 0% (clade A, among D. caudata sequences) to 5.2% (variation within clade E, represented by Diphtherophora sp. sequences from Iran; Table 3). On the other hand, genetic divergence between Diphtherophora clades was much higher, ranging from 8.2% (clade A vs. B) to as high as 23.7% (clade C vs. F). For comparison, genetic divergence between any of the Diphtherophora clades and Tylolaimophorus (also considered a representative of Diphtherophoridae) ranged from 25.6% (Tylolaimophorus vs. clade A) to 32.7% (Tylolaimophorus vs. clade F). Similarly, genetic divergence between Diphtherophora clades and trichodorid genera were in the same range (24.1% to 30.1%) as those observed for comparisons with Tylolaimophorus. This pattern is also consistent when comparing the genetic divergence between Diphtherophora clades and other outgroup taxa including genera Prismatolaimus de Man, 1880, Tripyla Bastian, 1865, and Tripylina Brzeski, 1963 (Table 3). 1 Taxon abbreviations: Tylolaimophorus (Tylo.), Trichodorus (Tric.), Paratrichodorus (Para.), Monotrichodorus (Mono.), Prismatolaimus (Prism.), Tripylina (Tripyli.). 2 Taxon represented by only one DNA sequence. With respect to the new species D. eldarica n. sp., genetic divergence ranged from 13.5% (between clade B, D. tenera + D. cf. obesa) to 19.5% (between clade E, Diphtherophora sp.). Although all six clades were strongly supported in our analyses, with ML branch support always greater than 98% and BPP always with maximum support, some relationships among Diphtherophora clades (i.e. some of the deeper nodes) were either weakly supported or not recovered by one of the inference methods (Fig. 4)., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on pages 379-382, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["de Man, J. G. (1880) Die Einheimischen, frei in der reinen Erde und im sussen Wasser lebende Nematoden [Free-living soil and freshwater nematodes]. Monographisch Bearbeitet. Tijdschrift der Nederlandsche dierk undige Vereniging, 5, 1 - 104. [in German]","Bastian, H. C. (1865) Monograph on the Anguillulidae. Transactions of the Linnean Society of London, 25, 73 - 184. https: // doi. org / 10.1111 / j. 1096 - 3642.1865. tb 00179. x","Brzeski, M. W. (1963) Nematode genera of the family Tripylidae (Nematoda, Enoplida). Acta Zoologica Cracoviensia, 8, 295 - 308."]}

Diphtherophora eldarica n. sp. (Figs. 1, 2, Table 1) Description. Female: Body habitus straight to slightly curved ventrally after fixation. Cuticle regularly striated from the anterior end to the level of secretory-excretory pore, but appearing smooth posteriorly; relatively thick, about one-fifth to two-fifths of the body width. Subcuticle layer finely striated along the entire body and about 2 ��m thick. Head region offset from the body contour. Outer labial papillae not readily observed. Amphid aperture oval in shape and as wide as about one-third of the head width at its base. Spear typical of the genus, with a sclerotized acute tip. Spear guiding apparatus arched, moderately sclerotized. Pharynx typical of the genus and composed of two parts, the anterior region (corpus) slender and expanding gradually to end as a rounded or sometimes slightly pyriform terminal bulb. Nerve ring located at mid-pharynx region or posterior to it. Secretory-excretory pore opposite the posterior end of the slender part of pharynx or at the beginning of the pharyngeal bulb. Cardia not observed under LM. Reproductive system didelphic, amphidelphic, with branches equally developed, ovaries reflexed with germinal zones directed toward vagina, spermatheca not present at the junction of oviduct and uterus. Sperm cells with rodshaped nuclei throughout the uteri. Vulva a transverse slit as seen in a ventral view. Cuticle invaginated just anterior and posterior to the vulva. Vagina orthogonal to the body axis, with moderately to well-developed musculature. Rectum 14���16 ��m or 54���73 % of the anal body width. Cuticle overhanging the anterior lip of the anus. Tail conoid, bent dorsally near tail tip, with a rounded terminus. The tail terminus lacks lateral caudal pores. Male: In general, morphologically similar to female. Body habitus almost straight. Cervical ventromedian papillae absent. No lateral body pores observed in the anterior region of the body. Reproductive system monorchic, extending to near mid-body, and containing numerous rod-shaped sperm. Two ventromedian precloacal supplements were observed, the anteriormost near the proximal end of spicules and the posterior one level with the distal ends of spicules. Spicules with marked capitulum and narrow shaft, expanding again at beginning of the blade part and then gradually tapering to the distal end. Spicular capsule weakly developed. Gubernaculum ventrally arcuate at anterior tip. Tail similar to female, with a rounded terminus. Overhanging cuticle on cloacal lips was not observed, as the cuticle did not separate from the body over a short distance anterior or posterior to cloaca. Diagnosis. The new species is characterized by females with striated cuticle at the anterior end of the body, head offset from the body contour, spear 15���17 ��m in length, rod-shaped sperm, overhanging cuticle on the anterior lip of the anus and a conical tail bent dorsally near the terminus. Additionally, males lack ventromedian neck papillae but have two ventromedian precloacal supplements at the level of the spicules. Type host and locality. The specimens were collected from the rhizosphere of pine trees (Pinus eldarica Medw.) in Dalampar valley in Margavar, Urmia city, West Azerbaijan province, Northwestern Iran by A. Mokaram Hesar in September 2015 (GPS coordinates: 37��10���65������N, 44��52���73������E). Type material. Holotype, three female paratypes and one male paratype, mounted in pure glycerin on a slide, were deposited in the collection of the Department of Plant Protection, School of Agriculture, Shiraz University, Shiraz, Iran. Etymology. The specific epithet refers to the associated plant species, Pinus eldarica, from the rhizosphere of which the new Diphtherophora species was recovered., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on pages 373-377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997

Diphtherophora perplexans (Van Der Linde, 1938) Goodey, 1951 (Fig. 3 C, G, Table 2) Morphologically, our populations from Iran fit well with the original population described from the USA (Cobb 1913). However, some morphometric characters also had larger ranges. Moreover, fine striations on the cuticle and large plate-like thickenings on the ventral or dorsal sides of the spear guiding apparatus were observed in some individuals, indicating close proximity of P. perplexans with P. pseudoperplexans (van der Linde, 1938) Goodey, 1951. In this study, four populations representing D. perplexans were found: one population was recovered from the rhizosphere of plum (Prunus domestica L.) trees in Mianeh, East Azerbaijan, Northwestern Iran; another population was recovered from the rhizosphere of apricot (Prunus armeniaca L.) trees in Khalkhal, Ardebil province, Northwestern Iran; two additional populations were recovered from turf grass in the campus of Shiraz University, at Eram Square, Shiraz, Fars province, Southern Iran., Published as part of Ghaderi, Reza, Hesar, Abas Mokaram, Karegar, Akbar & Pereira, Tiago Jos��, 2020, Molecular phylogeny of Diphtherophora de Man, 1880 (Nematoda: Diphtherophoridae), with description of a new species, pp. 371-385 in Zootaxa 4851 (2) on page 377, DOI: 10.11646/zootaxa.4851.2.10, http://zenodo.org/record/4476997, {"references":["Van der Linde, W. J. (1938) A contribution to the study of nematodes. Entomology Mem of Department of Agriculture and Forestry, Union of South Africa, 2, 1 - 40.","Goodey, T. (1951) Soil and freshwater nematodes. Methuen, London, 390 pp. https: // doi. org / 10.1097 / 00010694 - 195107000 - 00019","Cobb, N. A. (1913) New nematode genera found inhabiting freshwater and non-brackish soils. Journal of the Washington Academy of Sciences, 3, 432 - 444. https: // doi. org / 10.5962 / bhl. part. 20323"]}

Key to species of Cactodera (modified from Subbotin et al., 2010, Cid Del Prado Vera & Subbotin 2014) 1 Cyst generally two times or more longer than wide, mean L/W ratio = 2.3................................. C. estonica - Cyst usually less than twice as long as wide, mean L/W ratio = 1.1���1.8........................................... 2 2 Eggshell punctate...................................................................................... 3 - Eggshell smooth..................................................................................... 10 3 Mean stylet length of J2s �� 26 ��m......................................................................... 4 - Mean stylet length of J2s C. thornei - J2s tail length = 37���48 ��m, hyaline region = 17���24 ��m, fenestral diam. = 14���25 ��m........................ C. eremica 5 Mean J2s body length �� 411 ��m, mean tail length �� 42 ��m..................................................... 6 - Mean J2s body length C. milleri - Mean J2s tail length �� 43 ��m, b ratio> 3.5.................................................................. 7 7 Female L/W ratio C. cacti - Female L/W ratio �� 1.4, mean hyaline region of J2s �� 22 ��m................................... C. chenopodiae sp. n 8 Fenestral diam. C. galinsogae 9 Hyaline region of J2s = 4���8 ��m..................................................................... C. rosae - Hyaline region of J2s = 16���23 ��m.................................................................. C. evansi 10 Mean J2s tail length C. amaranthi - Mean J2s body length �� 406 ��m, mean hyaline region �� 16 ��m................................................. 12 12 Cyst with distinct vulval cone, J2s stylet length =21.0���23.0 ��m....................................... C. torreyanae - Cyst without distinct vulval cone, J2s stylet length =23.4���25.0 ��m......................................... C. salina 13 J2s stylet knobs anterior surface concave, DGO = 4.5���5.6 ��m................................................. 14 - J2s stylet knobs anterior surface convex, DGO = 2.5���3.0 ��m........................................... C. acnidae 14 Vulval denticles present, J2s tail length = 43���50 ��m..................................................... C. weissi - Vulval denticles absent, J2s tail length = 46���60 ��m................................................... C. radicale, Published as part of Feng, Yaxing, Wang, Dong, Xiao, Dongxue, Pereira, Tiago Jos��, Xuan, Yuanhu, Wang, Yuanyuan, Liu, Xiaoyu, Chen, Lijie, Duan, Yuxi & Zhu, Xiaofeng, 2018, Cactodera chenopodiae (Nematoda: Heteroderidae), a new species of cyst nematode parasitizing common lambsquarter (Chenopodium album) in Liaoning, China, pp. 361-375 in Zootaxa 4407 (3) on page 373, DOI: 10.11646/zootaxa.4407.3.4, http://zenodo.org/record/1216588, {"references":["Subbotin, S. A., Mundo-Ocampo, M. & Baldwin, J. G. (2010) Systematics of cyst nematodes (Nematodes: Heteroderinae). Vol. 8. Part A. Brill Academic Publishers, Leiden, 351 pp.","Cid Del Prado Vera, I. & Subbotin, S. A. (2014) A new cyst nematode, Cactodera torreyanae n. sp. (Tylenchida: Heteroderidae), parasitising romerito, Suaeda torreyana, in Texcoco, Mexico. Nematology, 16, 163 - 174. https: // doi. org / 10.1163 / 15685411 - 00002754"]}

Cactodera chenopodiae * n. sp. (Figs. 1���5, Tables 1) *specific epithet after the host, Chenopodium album L., Measurements. See Table 1. Description. Females. Body ovate to rounded or subspherical in shape with small vulval cone (Fig. 1C, 1D), female body pearly-white (Fig. 1G, 1H). Females with small vulval cones having slightly protruding lips (Fig. 1C, 1D). Female full of eggs and gelatinous egg sac not observed (Fig. 1I). Outer cuticular layer marked by a rugose pattern. Head slightly set off from the elongate and protruding neck, stylet and stylet knobs well developed (Fig. 1A). Excretory pore located at same level as end of isthmus (Fig. 1B). Anus distinct (Fig.1E, 1F). Cysts. Rounded to lemon-shaped, from light to dark brown with small vulval cone (Figs. 1K, 1L, 4A, 5F). Cyst surface with zigzag pattern at mid-body and not prominent on surface of the vulval cone (Fig. 4C, 4D). Cone is circumfenestrate and lacks an underbridge, bullae and vulval denticles (Fig. 1C, 1D, 4D). Anus distinct and encircled within a disc-like cuticular region (Fig. 4B). Males. Not found. J2s. Vermiform, tapering anteriorly and posteriorly (Fig. 2A). Stylet knobs rounded to slightly projecting anteriorly (Fig. 2B). Lip region slightly set off with four annuli. In en face view, an elongated labial disc surrounded by four submedial and two lateral lips (Fig. 4E, 5G). Excretory pore near level of gland lobe, hemizonid was one and a half annulus above excretory pore (Fig. 2B, 2E, 2F 5B). Lateral field with four lines with outer two ridges partially areolated along the body. (Figs. 2G, 4F, 5C). Tail tapering, with hyaline region; the hyaline region is often shorter than the stylet. Transition to hyaline region usually clearly demarcated by an outline that is V-shaped, U-shaped, or rarely sloping ventrally (Fig. 2C, 2D, 5D). Eggs. Surface with heavy punctations visible both under LM and SEM (Figs. 1J, 4G���I). Only found inside cysts. Type host and locality. Common lambsquarter, C. album in the Beiling Park, Shenyang, Liaoning province, China. Coordinates: 41��50���38��� N, 123��25���44��� E, 51 m a.s.l. Type material. Holotype female, seven paratype females, twenty paratype cysts and twenty paratype J2s were deposited for curation in the collection of the Nematology Institute of Northern China (NINC), Shenyang Agricultural University, Shenyang, China. Biology. Cactodera chenopodiae n. sp. was found on roots of C. album among other native vegetation of China. On the type host, the cyst nematode ranges from endoparasitic to semi-endoparasitic. Juveniles were detected on the roots of the host plant by acid fuchsin staining(Fig.3A). Moreover, some maturing juveniles (presumed to be sedentary J2s, J3s and J4s) were found with the anterior portion of their body penetrating into the roots (Fig. 3B���C). Diagnosis and relationships. Cactodera chenopodiae n. sp. belongs to the genus Cactodera which includes species characterized by a small vulval cone and a circumfenestrate terminal pattern. The new species can be differentiated from the fourteen other Cactodera species by a combination of morphological and molecular characters. The ranges of many morphological characters of C. chenopodiae overlap with those of other Cactodera species and isolates, including with certain C. cacti isolates from different regions of the US [e.g., Michigan, Graney & Bird (1990)]; yet, morphometric analyses of C. chenopodiae n. sp. demonstrate that the means for these morphological features are distinctive relative to C. cacti. For example, females of C. chenopodiae n. sp. have a larger L/W ratio [1.6 (1.4���1.7) vs. 1.2 (1.0���1.4)], and J2s have a smaller b ratio [3.8 (3.6���4.1) vs. 6 (5.6���6.8)] as well as a longer hyaline region [22.7 ��m (17.5���28.4) vs. 17.6 ��m (13.7���20.5)]. Cactodera chenopodiae n. sp. is also distinguished from C. milleri, known to parasitize common lambsquarter in Michigan, US (Graney & Bird, 1990), by the cyst having an anus set off within a distinctive disc-shaped cuticular pattern (Fig. 1E���F, 4B) and by having a greater fenestral diameter [23.5 ��m (19.9���26.3) vs. 18.7 ��m (14.3���22.0)]. Moreover, J2s stages have a larger b ratio [3.8 (3.6���4.1) vs. 2.9 (2.6���3.1)] and a longer hyaline region [22.7 ��m (17.5���28.4) vs. 18.2 ��m (14.6��� 21.2)] when compared to C. milleri. All measurements are in ��m. a CV: coefficient of variation. As for closely related species (i.e. based on molecular data), C. chenopodiae n. sp. is distinguished from C. estonica (Kirjanova & Krall, 1963) Krall & Krall, 1978, by the eggshell pattern (i.e., punctate in C. chenopodiae vs. smooth in C. estonica) and larger b ratio in J2s [3.8 (3.6���4.1) vs. 2.8 (2.7���2.9)]. In addition, C. chenopodiae n. sp. differs from C. rosae Cid del Prado & Miranda, 2008 by a longer hyaline region [22.7 ��m (17.5���28.4) vs. 6.3 ��m (4.0���6.8)], Published as part of Feng, Yaxing, Wang, Dong, Xiao, Dongxue, Pereira, Tiago Jos��, Xuan, Yuanhu, Wang, Yuanyuan, Liu, Xiaoyu, Chen, Lijie, Duan, Yuxi & Zhu, Xiaofeng, 2018, Cactodera chenopodiae (Nematoda: Heteroderidae), a new species of cyst nematode parasitizing common lambsquarter (Chenopodium album) in Liaoning, China, pp. 361-375 in Zootaxa 4407 (3) on pages 363-365, DOI: 10.11646/zootaxa.4407.3.4, http://zenodo.org/record/1216588, {"references":["Graney, L. S. O. & Bird, G. W. (1990) Descriptions and comparative morphology of Cactodera milleri n. sp. (Nematoda: Heteroderidae) and Cactodera cacti with a review and key to the genus Cactodera. Journal of Nematology, 22, 457 - 480.","Kirjanova, E. S. & Krall, E. (1963) Heterodera estonica n. sp. (Nematodes: Heteroderidae), Estonian cyst forming nematode. Izvestija Akademii Nauk Estonskoj SSR, Biologicheskaja Seria, 12, 219 - 223.","Krall, E. L. & Krall, K. A. (1978) Revision of the plant nematodes of the family Heteroderidae on the basis of the trophic specialization of these parasites and their co-evolution with their host plants. In: Fitogel'mintologicheskie Issledovaniya. USSR, Nauka, Moscow, pp. 39 - 56.","Cid Del Prado Vera, I. & Miranda, B. L. (2008) Second cyst-forming nematode parasite of barley (Hordeum vulgare L. Var. Esmeralda) from Mexico. Nematropica, 38, 105 - 114."]}