taxonID	type	description	language	source
03CD4B206627534C45AC92AEFC88A6C0.taxon	description	Figs. 1 – 3	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	description	Measurements. See Table 2.	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	materials_examined	Material examined. Type-material: Holotype cyst, three paratype vulval cones, one paratype male and five paratype J 2 s are deposited in the USDA Nematode Collection, Beltsville, Maryland; three paratype vulval cones, one paratype male and five paratype J 2 s in the Canadian National Nematode Collection, Ottawa, Canada. Other specimens are available in the Laboratory of Plant Nematology, South China Agricultural University, Guangzhou, China.	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	description	Description. Cyst. Lemon-shaped with distinct neck and vulval cone, light brown to dark brown in color. Stylet and other internal structures indistinct. Cuticle with irregular zig-zag pattern, but without remnant subcrystalline layer. Vulval cone ambifenestrate. Underbridge weak. Bullae absent. Eggs packed in the body. Anus distinct. White female. Body lemon-shaped, white in color, with prominent neck and vulval cone. Subcrystalline layer present. Head set off, with two annuli, the second annule larger and disc-shaped. Stylet long, with rounded knobs sloping slightly posteriorly. Dorsal pharyngeal gland duct opens 3 – 4 µm behind the basal knobs. Median bulb massive, with strongly developed valve plates. Excretory pore indistinct. Egg sac sometimes present, ranging from one-half to two times the size of the female, containing eggs. Male. Body vermiform, slightly curved ventrally when heat-relaxed. Lateral field with four incisures, outer bands areolated, wider than central band. Annulus width at mid-body about 1.8 µm. Head hemispherical, slightly offset, with a distinct labial disc and four or five postlabial distinct annuli. Stylet developed, with knobs slightly projecting or flat anteriorly. Dorsal pharyngeal gland duct opens 4 – 6 µm behind the basal knobs. Median bulb oval, with strong valve plates. Nerve ring slightly posterior to median bulb. Excretory pore 108 – 152.5 µm from anterior end of the body. Hemizonid 112 – 140 µm from anterior end of the body. Spicules arcuate, with finely bifurcate tip. Gubernaculum straight and simple. Tail very short and bluntly rounded. Phasmids indistinct. J 2. Body vermiform, slightly ventrally curved when heat-relaxed. Lateral field with three incisures. Body annuli 1.5 – 1.7 µm at mid-body. Head offset, hemispherical, with a labial disc and three or four indistinct postlabial annuli. Stylet robust with knobs slightly projecting or flat anteriorly. Dorsal pharyngeal gland duct opens 5.8 – 7 µm behind the basal knobs. Median bulb ovoid and prominent. Pharyngeal glands well developed. Hemizonid situated slightly anterior to excretory pore. Genital primordium oval, 8.0 (7.7 – 8.3) µm long, 5.8 (5.7 – 6.0) µm wide, 129.8 (100.7 – 143.7) µm anterior to tail terminus. Anus distinct. Tail conoid, narrowly tapering to a fine rounded terminus, hyaline terminal section distinct, 24.6 (18.3 – 33.8) µm long, occupying 50.1 (43.0 – 57.1) % of tail length. Phasmids pore-like, 5 – 8 annuli posterior to anus.	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	diagnosis	Diagnosis and relationships. Heterodera guangdongensis n. sp. belongs to the Cyperi group of Subbotin et al. (2010 a). Cysts are characterized by a protruding, ambifenestrate vulval cone with small fenestrae (22.0 – 33.8 µm in length), weak underbridge, short vulva-anus distance (28.9 – 35.9 µm) and a 31.1 – 41.0 µm long vulval slit, but without bullae; females are characterized by a 25.1 – 27.6 µm long stylet with rounded knobs sloping slightly posteriorly; males are characterized by a 21.5 – 23.0 µm long stylet with knobs slightly projecting or flat anteriorly, four incisures in lateral field, a 22.0 – 26.0 µm long spicule with bifurcate tip; J 2 s are characterized by a 19.3 – 21.3 long stylet with slightly projecting or anteriorly flattened knobs, three incisures in lateral field, a 41.7 – 61.3 µm long tail with finely rounded terminus and hyaline portion forming 50.1 (43.0 – 57.1) % of the tail length. In its morphology, the new species is most closely related to four members of the H. cardiolata Kirjanova & Ivanova, 1969 complex in the Cyperi group, including H. cardiolata, H. graminis Stynes, 1971, H. phragmitidis Kazachenko, 1986 and H. longicolla Golden & Dickerson, 1973. The new species differs from H. cardiolata by the more oblate fenestrae (22 – 33.8 µm vs 44 – 70 µm in length), the shorter vulva-anus distance (28.9 – 35.9 µm vs 42 – 50 µm), the longer female stylet (25.1 – 27.6 µm vs 19 – 21.4 µm) and the J 2 stylet knob shape (slightly projecting or anteriorly flattened vs prominently anteriorly directed). It differs from H. graminis by the longer female stylet (25.1 – 27.6 µm vs 19 – 24 µm), the smaller female DGO (3 – 4 µm vs 5 – 7 µm), the more oblate fenestrae (22.0 – 33.8 µm vs 32 – 63 µm in length), the shorter vulva-anus distance (28.9 – 35.9 µm vs 40 – 50 µm), the J 2 stylet knob shape (slightly projecting or anteriorly flattened vs prominent anteriorly concave), the male stylet knob shape (slightly projecting or flat anteriorly vs rounded) and the shorter spicules (22 – 26 µm vs 31 – 35 µm). From H. phragmitidis, it differs by the longer female stylet (25.1 – 27.6 µm vs 19.5 – 21 µm), the smaller female (223.6 – 398.5 × 207.9 – 367.9 µm vs 467 – 812 × 394 – 677 µm), the smaller cyst (266.5 – 415.1 × 207 – 361.2 µm vs 407 – 836 × 318.2 – 738 µm), the shorter spicules (22 – 26 µm vs 28.6 – 41.6 µm), the shorter gubernaculum (7.8 – 8 µm vs 11.7 – 14.3 µm), the male stylet knob shape (slightly projecting or flat anteriorly vs rounded) and the larger J 2 DGO (5.8 – 7 µm vs 3.6 – 4.8 µm). It can be distinguished from H. longicolla by the longer female stylet (25.1 – 27.6 µm vs 19 – 21 µm), the more oblate fenestrae (22.0 – 33.8 µm vs 37 µm in length), the longer male stylet (21.5 – 23 µm vs 17.9 – 20.2 µm), more incisures in the male lateral field (4 vs 3), more head annuli in the male (4 – 5 vs 3), the slightly shorter spicules (22 – 26 µm vs 25.2 – 28.6 µm), the shorter gubernaculum (7.8 – 8 µm vs 9 – 10.6 µm), the longer J 2 stylet (19.3 – 21.3 µm vs 16.8 – 18.5 µm) and the larger J 2 DGO (5.8 – 7 µm vs 3.4 – 6.1 µm).	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	discussion	The new species is also similar to other members of the Cyperi group, including H. cyperi Golden, Rau & Cobb, 1962, H. elachista Ohshima, 1974 and H. oryzicola Rao & Jayaprakash, 1978. It can be distinguished from H. cyperi by the longer female stylet (25.1 – 27.6 µm vs 22 – 22.4 µm), the smaller female DGO (3 – 4 µm vs 4.5 – 5.6 µm), the shorter female (223.6 – 398.5 µm vs 459 – 663 µm), the shorter cyst (266.5 – 415.1 µm vs 459 – 742 µm), the shorter males (624.0 – 861.3 µm vs 874 – 1159 µm), the shorter spicules (22 – 26 µm vs 28 – 30.8 µm), the shorter gubernaculum (7.8 – 8 µm vs 8 – 10 µm), the male stylet knob shape (slightly projecting or flat anteriorly vs rounded), the shorter J 2 (318.1 – 409.5 µm vs 414 – 532 µm), the slightly larger J 2 DGO (5.8 – 7 µm vs 3.8 – 6.1 µm) and the slightly shorter J 2 tail (41.7 – 61.3 µm vs 56 – 79 µm); from H. elachista by the longer female stylet (25.1 – 27.6 µm vs 19 – 23 µm), the absence of bullae (vs presence), the absence of subcrystalline in cyst stage (vs presence), the longer male stylet (21.5 – 23 µm vs 20 – 21 µm), the shorter spicules (22 – 26 µm vs 26 – 29 µm), the shorter gubernaculum (7.8 – 8 µm vs 8 – 13 µm) and the male stylet knob shape (slightly projecting or flat anteriorly vs sloping posteriorly); from H. oryzicola by the shorter female (223.6 – 398.5 µm vs 414 – 520 µm), the longer female stylet (25.1 – 27.6 µm vs 18 – 20 µm), the absence of bullae (vs presence), the shorter males (624 – 861.3 µm vs 896 – 980 µm) and the J 2 stylet knob shape (slightly projecting or anteriorly flattened vs rounded). Compared with the four Heterodera species from bamboo, i. e. H. bamboosi, H. koreana, H. hainanensis and H. fengi, H. guangdongensis n. sp. can be easily distinguished from the first three species by the presence of fenestrae (vs afenestrate). In addition, the new species further differs from H. bamboosi by the presence of a prominent vulval cone (vs absence of vulval cone) and more incisures in the male lateral field (4 vs 3); from H. koreana and H. hainanensis by more J 2 lip annuli (3 – 4 vs 2). H. guangdongensis n. sp. differs from H. fengi by the longer female stylet (25.1 – 27.6 µm vs 23 – 24 µm), the smaller fenestrae (22.0 – 33.8 × 27.8 – 35.5 µm vs 40 – 65 × 45 – 62.5 µm), the shorter vulval slit (31.1 – 41 µm vs 40 – 60 µm), the shorter male stylet (21.5 – 23 µm vs 24.5 – 26.3 µm), the male stylet knob shape (slightly projecting or flat anteriorly vs rounded, sloping posteriorly), the shorter spicules (22 – 26 µm vs 27.5 – 31.3 µm), the shorter J 2 (318.1 – 409.5 µm vs 440 – 520 µm), the shorter J 2 stylet (19.3 – 21.3 µm vs 22 – 24 µm), the larger J 2 DGO (5.8 – 7 µm vs 4.5 – 5.3 µm), the shorter J 2 tail (41.7 – 61.3 µm vs 62.5 – 77 µm) and the shorter J 2 hyaline tail portion (18.3 – 33.8 µm vs 35 – 45 µm).	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	materials_examined	Type-locality and habitat: Heterodera guangdongensis n. sp. was originally collected from roots and soils around roots of bamboo (Phyllostachys pubescens) in Guangzhou City (latitude N 23 ° 9 ' 23 '', longitude E 113 ° 21 ' 40 ''), Guangdong Province, China in 2009. The soil type is sandy loam and the local climate is subtropical. Other locality: Other than the type locality, H. guangdongensis n. sp. was also collected from roots and soils around roots of P. pubescens in Nanning City (latitude N 22 ° 52 ' 44 '', longitude E 108 ° 25 ' 3 '') and Hechi City (latitude N 24 ° 33 ' 29 '', longitude E 107 ° 46 ' 30 ''), Guangxi Province, China in 2011. The soil type is sandy loam and the local climate is also subtropical.	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	etymology	Etymology: The specific name refers to the type locality, Guangdong Province.	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
03CD4B206627534C45AC92AEFC88A6C0.taxon	description	Molecular profiles and phylogenetic status. Three rDNA amplicons of 784 - bp LSU D 2 D 3 were sequenced, obtained from single J 2 s collected from Guangzhou, Hechi and Nanning, respectively. GenBank accession numbers of the three sequences are JX 081320, KM 224882 and JX 081321. The identities among these three D 2 D 3 sequences of H. guangdongensis n. sp. were 99.6 – 99.7 %. A BlastN search of H. guangdongensis n. sp. on the LSU D 2 D 3 showed high-scoring matches with some Heterodera species, the highest being with two sequences of H. elachista (GenBank accession number HM 560842 and HM 560843). The identities of these two sequences and the three D 2 D 3 sequences from the new species ranged from 96.8 % to 97.2 %. Three ITS-rDNA sequences, two 1161 bp and one 1163 bp, were obtained, respectively, from the same J 2 DNA templates mentioned above. GenBank accession numbers of these three sequences are JX 081324, KM 224880 and KM 224881. The identities among these three ITS sequences of H. guangdongensis n. sp. were 99.7 %. A BlastN search of H. guangdongensis n. sp. on the ITS also indicated high-scoring matches with some Heterodera species, but the identities were less than 90 %. The highest match was also two sequences of H. elachista (GenBank accession number JN 201917 and JN 201914) with 87.5 % to 88 % identities. The molecular phylogenetic relationships of H. guangdongensis n. sp. reconstructed by our analyses are presented in Figures 4 and 5. A phylogenetic tree based on LSU D 2 D 3 from a multiple alignment of 676 total characters was constructed (Figure 4). The average nucleotide composition is as follows: 16.98 % A, 22.45 % C, 34.34 % G and 26.23 % T. When Cryphodera brinkmani Karssen & van Aelst, 1999 was used as the outgroup taxon, all Heterodera species form a monophyletic group with strong support [100 % posterior probability (PP)]. H. guangdongensis n. sp. is positioned in a 100 % - supported clade with several species in the Cyperi group, such as H. elachista, H. fengi and H. oryzicola. A phylogenetic tree based on ITS-rRNA was generated from a multiple alignment of 1174 total characters (Figure 5). The average nucleotide composition is as follows: 17.6 % A, 23.15 % C, 30. 25 % G and 29.01 % T. When using H. circeae Subbotin & Sturhan, 2004 as an outgroup taxon, H. guangdongensis n. sp. is sister to H. elachista in the Cyperi group, with high support (94 % PP), and they were placed in a moderately supported clade (82 % PP) with other species including H. oryzicola, H. cyperi and H. mothi belonging to the Cyperi group. This clade is closely related to another member in the Cyperi group, H. fengi, with 100 % support. The RFLP-ITS-rRNA profile of H. guangdongensis n. sp. is presented in Figure 6 and Table 3, and differs from profiles of other known Heterodera species, as seen in Subbotin et al. (2010 a), especially from profiles of those species in the Cyperi group, including H. cardiolata, H. cyperi, H. elachista, H. fengi, H. mothi Khan & Husain, 1965 and H. oryzicola, obtained with the same PCR primers and the same set of restriction enzymes, as summarized by Subbotin et al. (2010 b) and Wang et al. (2013).	en	Zhuo, Kan, Wang, Honghong, Zhang, Hongling, Liao, Jinling (2014): Heterodera guangdongensis n. sp. (Nematoda: Heteroderinae) from bamboo in Guangdong Province, China-a new cyst nematode in the Cyperi group. Zootaxa 3881 (5): 488-500, DOI: 10.11646/zootaxa.3881.5.4
