Camallanus sodwanaensis, Svitin & Truter & Kudlai & Smit & Preez, 2019

Svitin, Roman, Truter, Marliese, Kudlai, Olena, Smit, Nico J. & Preez, Louis du, 2019, Novel information on the morphology, phylogeny and distribution of camallanid nematodes from marine and freshwater hosts in South Africa, including the description of Camallanus sodwanaensis n. sp., International Journal for Parasitology: Parasites and Wildlife 10, pp. 263-273 : 265-270

Genus Camallanus Railliet et Henry, 1915 .

Camallanus sodwanaensis n. sp.

Type host: Dusky sweeper Pempheris adusta Bleeker, 1877 ( Perciformes : Pempheridae ).

Other hosts: Stocky hawkfish Cirrhitus pinnulatus (Förster, 1801) ( Perciformes : Cirrhitidae ), Banded grunter Pomadasys furcatus (Bloch et Schneider, 1801) ( Perciformes : Haemulidae ), Jarbua terapon Terapon jarbua (Forsskål, 1775) ( Perciformes : Terapontidae ), and Largespotted dart Trachinotus botla (Shaw, 1803) ( Perciformes : Carangidae ).

Site of infection: Intestine.

Type locality: Sodwana Bay , KwaZulu-Natal Province, South Africa (32̊40′46"E; 27̊32′24"S).

Type material: Holotype (male, [ NMB P 509]), allotype (female, [ NMB P 510]), paratypes [ NMB P 511] deposited in the National Museum Parasite Collection (Bloemfontein, South Africa).

Intensity: Pempheris adusta : 1–15 (5.7); Cirrhitus pinnulatus : 1–5 (2.3); Terapon jarbua : 1–2 (1.5); Pomadasys furcatum : 1–1 (1); Trachinotus botla : 1; total: 1–15 (3.0).

Prevalence: Pempheris adusta – 14% (six of 22 specimens were infected); Cirrhitus pinnulatus – 75% (three of four specimens were infected); Terapon jarbua – 67%; Pomadasys furcatum – 50% (two of four specimens were infected); Trachinotus botla – 25% (one of four specimens were infected); total – 27%.

Abundance: Pempheris adusta – 0.8; Cirrhitus pinnulatus – 1.8; Terapon jarbua – 1; Pomadasys furcatum – 0.5; Trachinotus botla – 0.3; total – 0.8.

Representative DNA sequences: 28S [ MN 525306], 18S [ MN 514774].

ZooBank registration: To comply with the regulations set out in article 8.5 of the amended 2012 version of the International Code of Zoological Nomenclature ( ICZN, 2012), details of the new species have been submitted to ZooBank. The Life Science Identifier ( LSID) for Camallanus sodwanaensis n. sp. is urn:lsid:zoobank.org:act:BCEC589F-46B3-4645-9EC1-F6FA36B11213

Etymology: The species is named after its type locality.

Description ( Figs. 1 and 2 ).

General. Body thin, elongated with maximum width at mid-length. Females generally larger than males. Cuticle with conspicuous transverse and fine longitudinal striations. Apical: oral opening narrow slit-like, surrounded by four conspicuous cephalic papillae ( Fig. 1C ; 2F ). Four sclerotised plates situated on external surface of buccal capsule valves near their anterior margin. Buccal capsule with well developed valves supported by numerous ridges ( Fig. 1A,B,C ; 2C,D,F,G ). Thick sclerotised basal ring present at base of buccal capsule. Oesophageal cup well developed. Two prominent tridents situated on ventral and dorsal sides of buccal capsule valves. Dorsal and ventral tridents equal in size and shape, each consisted of three posteriorly directed prongs. Central prong somewhat longer than sublateral ones, often reaching anterior margin of nerve ring ( Fig. 1A,B,E ; 2B,D,E ). Muscular and glandular oesophagus almost cylindrical, slightly widening in posterior third. Nerve ring encircling oesophagus close to its anterior end. Excretory pore situated at level of nerve ring or slightly posterior to it ( Fig. 1A ; 2B ). Deirids not observed. Intestine and rectum straight, narrow. Tail tapering with prominent phasmids situated at level of its anterior third.

Males. Measurements based on three specimens. Body 4.7–8.5 (6.6) [8.5] mm long, 112–173 (152) [173] maximum wide ( Fig. 2A ). Buccal capsule valves 91–101 (95) [91] long, 84–95 (92) [87] maximum wide, supported by 23–25 (25) [23] ridges, of which 9–9 (9) [9] incomplete. Basal ring 16–27 (22) [22] long, 61–69 (66) [69] wide. Oesophageal cup 13–27 (19) [16] long, 25–32 (30) [32] wide. Dorsal trident 134–160 (147) [160] long, 20–25 (23) [23] wide in lateral projection, ventral trident 131–169 (148) [160] long, 24–25 (25) [25] wide.

Muscular oesophagus 655–880 (794) [880] long, 10–14 (12) [10]% of body length; 62–79 (71) [62], 82–98 (90) [98] and 100–113 (107)

[113] wide at anterior, mid-length and posterior levels, respectively. Glandular oesophagus 562–940 (767) [940] long, 11–12 (12) [12]% of body length; 67–89 (78) [89], 77–88 (83) [88] and 82–110 (96) [110] wide at anterior, mid-length and posterior levels, respectively. Nerve ring at 205–211 (207) [211], 24–31 (27) [24]% of muscular oesophagus length. Excretory pore at 210–242 (218) [242] from anterior end of body, 3–4 (4) [3]% of body length.

Caudal alae narrow, supported by papillae: eight pairs of pedunculated precloacal; two pairs of adcloacal (anterior and posterior to cloaca), five pairs of postcloacal (two pairs grouped slightly posterior to cloaca, one pair at level of tail mid-length and one close to tail end) ( Fig. 1D ; 2I ). Spicules unequal, simple-shaped with sharpened tips ( Fig. 1G ; 2H ). Right spicule prominent, 303–328 (313) [303] long, 4–7 (5) [4]% of body length; left one less sclerotised, poorly visible 162–205 (184) [205] long, 2–3 (3) [2]% of body length. Tail conical, tapering to rounded tip, 86–101 (94) [94] long, 1–2 (2) [2]% of body length.

Females. Measurements based on nine gravid (larvigerous) specimens. Body 6.0–11.8 (9.0) [10.9] mm long, 130–283 (222) [316] wide. Buccal capsule valves 106–173 (141) [115] long, 102–178 (141) [111] wide, supported by 19–33 (28) [28] ridges, of which 12–20 (16) [14] incomplete. Basal ring 20–30 (25) [22] long, 64–99 (83) [68] wide. Oesophageal cup 14–26 (21) [23] long, 30–40 (35) [35] wide. Dorsal trident 136–205 (171) [152] long, 20–36 (28) [20] wide in lateral projection, ventral one 134–206 (173) [151] long, 17–34 (28) [21] wide.

Muscular oesophagus 800–1173 (1053) [1087] long, 10–14 (12) [10]% of body length; 75–98 (88) [68], 89–124 (108) [112] and 104–154 (132) [113] wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 758–1152 (938) [1141] long, 9–14 (11) [10]% of body length; 86–126 (106) [63], 78–134 (111) [81] and 100–140 (123) [103] wide at anterior, mid-length and posterior level, respectively. Nerve ring at 202–287 (252) [246] from anterior end of body, 17–29 (24) [23]% of muscular oesophagus length. Excretory pore at 210–308 (263) [275], 2–4 (3) [3]% of body length. Viviparous. Vulva with distinct lips ( Fig. 2J ), opening posterior to small projection to body wall at 3.5–5.7 (4.8) [5.1] mm from anterior end of body, 46–58 (53) [46]% of body length. Tail 87–140 (104) [111] long, 1–2 (1) [1]% of body length. Tail tip rounded in mature females and bearing two small mucrons in immature ones ( Fig. 1F ; 2K ).

Remarks. The species belongs to the genus Camallanus based on the presence of a well developed buccal capsule consisting of two valves, each supported by longitudinal ridges (not divided in dorsal and ventral group with a gap between), and presence of tridents on the dorsal and ventral sides of the buccal capsule valves ( Anderson et al., 2009). Camallanus sodwanaensis n. sp. is the first species of the genus found in marine fish from Southern Africa. Only two species of Camallanus described from African freshwater fishes are still considered as valid and were subsequently found after the first description: C. longicaudatus and C. kirandensis . The new species can be distinguished from C. longicaudatus by the relatively smaller length of the female tail (1–2% of body length vs 12–14%) and number of postcloacal papillae in males (6

in C. longicaudatus vs 5 in C. sodwanaensis n. sp.) ( Moravec, 1973). By the same characters, C. sodwanaensis n. sp. can be easily distinguised from C. kirandensis that has a comparatively long tail (868–1400 long in 8.4–20.0 mm long females, comprising approximately 7%). and only three pairs of postcloacal papillae ( Baylis, 1928; Amin, 1978). Out of the species described from marine fish, C. sodwanaensis n. sp. is morphologically (size and shape of buccal capsule and tridents, morphology male spicules, general body measurements) and geographically the most closely related to C. carangis . The most reliable character to distinguish between the two species is the number of postcloacal papillae –

five pairs in C. sodwanaensis n. sp. and six pairs in C. carangis ( Rigby et al., 1998) .

Genus Paracamallanus Yorke et Maplestone, 1926 .

Paracamallanus cyathopharynx (Baylis, 1923) .

Host: African sharptooth catfish Clarias gariepinus (Burchell, 1822) .

Locality: Ndumo Game Reserve, KwaZulu-Natal Province, South Africa (32̊30′69"E; 26̊85′63"S).

Site of infection: Intestine.

Intensity: 1–8 (3.3).

Prevalence: 46% (seven of 15 infected).

Abundance: 1.5.

Representative DNA sequences: 18S [MN514775], COI [MN523683].

Description ( Fig. 3 ).

General. Medium-sized nematode, body thin with maximum width at mid-length. Cuticle with conspicuous transverse striations along entire body. Apical: oral opening slit-like, surrounded by four cephalic plates, four conspicuous outer cephalic papillae, 4 min inner cephalic papillae and two amphids ( Fig. 3C ). Buccal capsule well sclerotised, divided in anterior and posterior parts. Anterior part consisting of two valves, each supported by nine longitudinal ridges and two tridents on dorsal and ventral sides ( Fig. 3B,E ). Each trident consisted of three posteriorly directed prongs of which central one somewhat longer than sublateral. Dorsal and ventral tridents equal in size and shape, beginning at level of buccal capsule anterior quarter and ending at level of oesophageal cup. Posterior part of buccal capsule shorter and narrower than anterior one with thick well-sclerotised walls. Oesophageal cup shorter than wide, poorly sclerotised. Muscular oesophagus evenly widened from anterior to posterior part. Glandular oesophagus almost cylindrical, slightly widened in middle third. Nerve ring encircling muscular oesophagus at level of its anterior third. Excretory pore opening somewhat posterior to level of nerve ring ( Fig. 3A ). Intestine and rectum straight, narrow. Tail tapering.

Males. Measurements based on nine specimens. Body 1.6–6.8 (5.4) mm long, 46–124 (102) wide. Anterior part of buccal capsule 51–61 (58) long, 53–58 (55) wide. Posterior part of buccal capsule 35–41 (38) long, 38–53 (46) wide. Oesophageal cup 4–8 (6) long, 8–18 (13) wide. Dorsal trident 65–76 (71) long, 9–12 (11) wide in lateral projection, ventral one 65–77 (71) long, 9–13 (11) wide. Muscular oesophagus 204–469 (384) long, 5.8–12.8 (7.5)% of body length; 31–47 (41), 30–62 (47) and 37–73 (57) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 210–670 (531) long, 7.8–13.2 (10.2)% of body length; 33–63 (52), 44–72 (56) and 44–73 (54) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 134–165 (147) from anterior end, 32.0–65.7 (40.0)% of muscular oesophagus. Excretory pore at 149–251 (196) from anterior end, 2.6–4.9 (3.4)% of body length.

Posterior end coiled ventrally. Caudal alae narrow, supported by papillae: five pairs of precloacal pedunculated papillae, two pairs of adcloacal papillae (anterior and posterior to cloaca), six pairs of postcloacal papillae (three pairs grouped posterior to cloaca, two pairs at mid-length of tail, one pair close to tail end). Spicules unequal. Right one longer, well-sclerotised, 177–271 (223) bearing short process on its tip, 27–69 (43) long. Left spicule shorter, less sclerotised, simple-shaped with sharpened tip, 37–70 (52) long. Tail tapering with rounded tip, 59–71 (66) long ( Fig. 3H ).

Females. Measurements based on seven gravid specimens. Body 9.5–15.8 (11.7) mm long, 130–204 (167) wide ( Fig. 3D ). Anterior part of buccal capsule 71–84 (76) long, 67–84 (75) wide. Posterior part of buccal capsule 48–56 (53) long, 63–70 (66) wide. Oesophageal cup 7–10 (9) long, 12–22 (17) wide. Dorsal trident 80–112 (93) long, 11–15 (13) wide in lateral projection, ventral one 78–112 (92) long, 11–16 (14) wide. Muscular oesophagus 535–681 (581) long, 3.5–5.8 (5.1)% of body length; 48–66 (58), 54–74 (68) and 70–110 (87) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 680–955 (793) long, 6.0–7.4 (6.8)% of body length; 61–91 (71), 70–92 (80) and 76–119 (92) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 173–213 (192) from anterior end, 28.8–38.4 (33.2)% of muscular oesophagus. Excretory pore at 227–420 (275) from anterior end, 1.6–4.2 (2.4)% of body length.

Vulva with slightly elevated lips at 3.7–8.5 (6.1) mm from anterior end, 37.1–56.3 (52.1)% of body length ( Fig. 3F ). Tail tapering, 291–507 (362) long, bearing three small mucrons on tip ( Fig. 3G ).

Remarks. The species has been found in many localities throughout Africa from clariid catfishes Mwita (2011); Madanire-Moyo and Barson (2010); Ajala and Fawole (2014); Moravec and Van As (2015c); Moravec and Jirků (2017) and was reported once from Israel ( Paperna, 1964). Nevertheless, the morphology of the species was illuminated only in the latest redescription provided by Moravec and Van As (2015c). In the redescription the authors described an unusual shape of the right spicule consisting of two parts: thin elongated anterior and short well sclerotised posterior that has often been confused with the gubernaculum or the left spicule. At the same time, the left spicule was described as poorly sclerotised and needle-like. In present study, we also found a clearly visible right spicule consisting of two parts and poorly sclerotised (visible only on high magnification with DIC and when dissected) left one, both with slightly wider ranges of measurement values. Also, similar to that in the latest redescription, we found eight cephalic papillae on the anterior end of nematodes. Despite that papillae of inner circle are minute and often covered with host tissue, they were clearly observed under the light microscope using high magnification and DIC.

Rigby and Rigby (2014) proposed Paracamallanus as a junior synonym of the genus Oncophora based on the similarities in their buccal capsule morphology. These authors suggested that the only difference between genera is the greater width of the female posterior to the vulva in Oncophora and assumed it as not indicative of different genera. In our opinion, significance of the characters for generic differentiation should be confirmed with sufficient molecular analyses. Therefore, in the present study we prefer to assign found species to genus Paracamallanus following Moravec and Van As (2015c) and Moravec and Scholtz (2017).

Genus Procamallanus Baylis, 1923 .

Procamallanus pseudolaeviconchus Moravec et Van As, 2015 .

Host: African sharptooth catfish Clarias gariepinus (Burchell, 1822) .

Locality: Ndumo Game Reserve, KwaZulu-Natal Province, South Africa (32̊30′69"E; 26̊85′63"S).

Site of infection: Intestine.

Intensity: 1–2 (1.3).

Prevalence: 46% (seven of 15 infected).

Abundance: 0.6.

Representative DNA sequences: 18S [MN514770], 28S [MN525307], COI [MN523682].

Description ( Fig. 4 ).

General. Body thin, elongated with maximum width at mid-body region. Cuticle with prominent transverse striations. Apical: oral opening rounded with unlobed peribuccal flange, surrounded by four inner submedian papillae, four outer submedian papillae and two amphids on lateral sides ( Fig. 4C ). Buccal capsule well sclerotised, longer than wide with two step-like folds and wide basal ring on its base ( Fig. 4D ). Oesophageal cup small, poorly sclerotised. Muscular oesophagus club-shaped with elongated posterior bulb. Glandular oesophagus almost two times longer than muscular one, almost cylindrical slightly widened posteriorly. Nerve ring encircling muscular oesophagus somewhat anterior to its mid-length. Excretory pore opening at level of muscular oesophagus posterior quarter ( Fig. 4B ). Minute deirids situated posterior to level of nerve ring. Intestine straight, narrow. Rectum straight, with thin walls. Tail tapering with rounded tip in both sexes.

Males. Measurements based on three specimens. Body 5.1–5.9 (5.5) mm long, 114–118 (116) wide ( Fig. 4A ). Buccal capsule 53–61 (56) long, 38–39 (39) wide. Basal ring 7–9 (8) long, 25–26 (26) wide. Oesophageal cup 7–7 (7) long, 10–12 (11) wide. Muscular oesophagus 365–388 (373) long, 6.5–7.2 (6.8)% of body length; 29–33 (31), 38–41 (40) and 48–58 (54) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 667–824 (740) long, 13.1–13.9 (13.4)% of body length; 44–47 (45), 54–64 (58) and 51–62 (55) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 173–189 (181) from anterior end of body, 47.4–49.2 (48.4)% of muscular oesophagus length. Excretory pore at 221–384 (326) from anterior end of body, 4.3–6.7 (5.8)% of body length.

Posterior end coiled ventrally with narrow caudal alae supported by papillae: nine pairs of precloacal pedunculated papillae, one pair of adcloacal papillae (anterior to cloaca) and four pairs of postcloacal papillae ( Fig. 4H ). Spicules unequal, simple-shaped with sharply pointed distal ends. Right spicule clearly visible, 112–126 (118) long; left one less sclerotised, 42–47 (45) long. Gubernaculum poorly sclerotised, 43 long (measured in one specimen). Tail tapering with rounded tip 46–54 (51) long.

Females. Measurements based on three gravid species. Body 4.9–9.1 (7.6) mm long, 110–186 (150) maximum width ( Fig. 4E ). Buccal capsule 58–67 (61) long, 54–56 (55) wide. Basal ring 9–10 (9) long, 26–35 (31) wide. Oesophageal cup 11–11 (11) long, 14–22 (18) wide. Muscular oesophagus 418–470 (448) long, 5.0–8.6 (6.3)% of body length; 27–37 (34), 32–56 (45) and 53–71 (62) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 577–814 (711) long, 8.5–11.8 (9.8)% of body length; 37–58 (50), 53–62 (59) and 49–61 (57) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 201–217 (209) from anterior end, 45.7–48.1 (46.7)% of muscular oesophagus. Excretory pore at 261–367 (315) from anterior end, 3.0–6.5 (4.5)% of body length. Vulva postequatorial, opening posterior to small projection of body wall at 3.0–6.5 (4.5) mm from anterior end, 58–63 (60)% of body length ( Fig. 4G ). Tail tapering with rounded tip, 89–114 (98) long, 1.0–1.8 (1.4)% of body length ( Fig. 4F ).

Remarks. The species was recently described by Moravec and Van As (2015a) based on material collected from the catfish Cl. gariepinus from Egypt and Botswana. The morphology and measurements of the specimens reported here from South Africa generally correspond with the original decription and the specimens represent a new geographical record.

Barson and Avenant-Oldewage (2006) reported P. leaviconchus from Cl. gariepinus in South Africa. Although, based on the provided SEM images it is clear that the peribuccal flange of the parasites is rounded, corresponding to that of P. pseudolaeviconchus (contrary to six-lobed in P. leaviconchus ).

Genus Spirocamallanus Olsen, 1952 .

Spirocamallanus daleneae ( Boomker, 1993) .

Host: Brown squeaker Synodontis zambezensis (Peters, 1852) .

Locality: Ndumo Game Reserve, KwaZulu-Natal Province, South Africa (32̊30′69"E; 26̊85′63"S).

Site of infection: Intestine.

Intensity: 1–2 (1.6).

Prevalence: 32% (eight of 25 infected).

Abundance: 0.52.

Representative DNA sequences: 28S [MN525304], 18S [MN514771].

Description ( Fig. 5 ).

General. Comparatively long nematodes, body thin with maximum width at mid-length. Cuticle with conspicuous transverse striations along entire body. Apical: oral opening rounded surrounded by 6 min papillae, four inner submedian papillae, four outer submedian papillae and two amphids ( Fig. 5B ). Buccal capsule sclerotised, longer than wide, with 9–14 (of which anterior and posterior ones usually incomplete) spiral ridges ( Fig. 5E ). Basal ring short and narrow, oesophageal cup poorly developed. Buccal capsule supported by six columns each consisting of four blocks ( Fig. 5D,F ). Muscular oesophgus club-shaped, almost cylindrical in anterior half with elongated posterior bulb. Glandular oesophagus somewhat shorter than muscular one, almost cylindrical along whole length, slightly widening posteriorly. Nerve ring encircling muscular oesophagus at level of its mid-length. Position of excretory pore varying within level of muscular oesophagus posterior quarter. Intestine and rectum strait, narrow. Tail tapering without mucrons.

Males. Measurements based on six specimens. Body 1.6–2.0 (1.8) mm long, 246–354 (295) wide ( Fig. 5A ). Buccal capsule 83–107 (95) long, 77–95 (88) wide with 11–14 (13) ridges. Basal ring 9–14 (11) long, 43–55 (51) wide. Muscular oesophagus 704–737 (717) long, 3.6–4.4 (4.1)% of body length; 54–69 (64), 51–72 (62) and 84–126 (104) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 588–690 (628) long, 3.3–3.6 (3.5)% of body length; 65–91 (82), 91–129 (107) and 77–108 (96) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 348–427 (373) from anterior end, 48.6–57.9 (52.1)% of muscular oesophagus length. Excretory pore opening at 473–652 (565) from anterior end, 2.4–3.7 (3.1)% of body length. Caudal end bended ventrally with narrow caudal alae supported by papillae: three pairs pre-anal pedunculated papillae, two pairs of sessile ad-anal papillae (one anterior and one posterior to cloaca), four pairs of post-cloacal papillae (of which posterior one situated close to alae margins) ( Fig. 5G ). Spicules unequal, poorly sclerotised. Right spicule larger, with bifurcated tip (with one branch somewhat longer), 163–231 (205) long; left one shorter, simple-shaped with sharpened tip, 132–199 (164) long. Tail 212–271 (235), 1.0–1.3 (1.1)% of body length.

Females. Measurements based on six gravid specimens. Body 1.2–3.0 (2.1) mm long, 227–589 (383) wide ( Fig. 5C ). Buccal capsule 59–123 (97) long, 63–118 (93) wide, supported by 9–13 (11) ridges. Basal ring 8–14 (10) long, 40–68 (55) wide. Muscular oesophagus 605–950 (798) long, 3.2–5.4 (4.2)% of body length; 49–78 (63), 52–77

(66) and 71–142 (107) wide at anterior, mid-length and posterior level, respectively. Glandular oesophagus 427–927 (670) long, 2.8–4.0 (3.4)% of body length; 68–101 (84), 72–136 (105) and 80–113 (97) wide at anterior, mid-length and posterior level, respectively. Nerve ring at 271–479 (348) from anterior end, 37.6–50.4 (43.8)% of muscular oesophagus length. Excretory pore at 392–766 (324) from anterior end, 2.3–4.6 (3.1)% of body length. Vulva small, often poorly visible, opening around mid-body level at 5.5–17.4 (10.4) mm from anterior end, 46.9–64.3 (53.7)% of body length ( Fig. 5I ). Tail conical, bearing short process with rounded tip ( Fig. 5H ).

Remarks. The morphology of the specimens collected from the Ndumo Game Reserve corresponds to the original description of S. daleneae from the Brown squeaker Sy. zambezensis collected in South Africa's Kruger National Park ( Boomker, 1993). The only difference found is that our specimens have eight columns around the buccal capsule which are not reported (probably overlooked) in the original description. Nevertheless, all other morphological (number of buccal capsule ridges, shape of tail in females, number and arrangement of papillae on male caudal region) and morphometric data, as well as host species and geographical origin, led us to assign found specimens to S. daleneae . Outside South Africa, S. daleneae has been recorded from Sy. acanthomias Boulenger, 1899 in the Central African Republic (Moravec and Jirků, 2015b) and from Sy. vanderwaali Skelton et White, 1990 in Botswana ( Moravec and Van As, 2015b). The authors assigned the studied specimens to S. daleneae , but mentioned that in their material all specimens possessed a nerve ring more anterior than that in the original description ( Boomker, 1993). Moreover, Moravec and Jirků (2015) described five pairs of postcloacal papillae in males contrary to the four pairs presented in the original description. These authors assumed that Boomker (1993) overlooked one pair of caudal papillae in male and the nerve ring position. Nevertheless, all specimens in our material from the type host of S. daleneae Sy. Zambezensis , from South Africa possessed a nerve encircling muscular oesophagus posterior to its mid-length and all males possessed four pairs of postcloacal papillae. In our opinion, the specimens studoed by Moravec and Van As (2015b) and Moravec and Jirků (2015) might belong to a new species while S. daleneae might be a specific parasite of Sy. zambezensis .

Moravec and Jirků (2015) and Moravec and Van As (2015b) assigned the species to the genus Procamallanus and subgenus Spirocamallanus . In the present study, we prefer to assign the species to Spirocamallanus as a separate genus due to distant phylogenetic relationships between S. daleneae and P. pseudolaeviconchus (24% (189 nt) in the 28S rDNA gene) (see Table 1).

Batrachocamallanus xenopodis (Baylis, 1929) .

Host: Muller's platanna Xenopus muelleri (Peters, 1844) .

Locality: Ndumo Game Reserve, KwaZulu-Natal Province, South Africa (32̊32′34"E; 26̊93′11"S).

Site of infection: Stomach.

Intensity: 1–4 (2.7).

Prevalence: 100% (three of three infected).

Abundance: 2.7.

Representative DNA sequences: 28S [MN525305], 18S [MN514768], COI [MN523681].

Description ( Fig. 6 ).

General. Small nematodes, body comparatively thick with maximum width at anterior quarter. Cuticle with conspicuous transverse striations along entire body. Apical: oral opening rounded surrounded by 6 min papillae, four inner submedian papillae, four outer submedian papillae and two amphids ( Fig. 6B ). Buccal capsule sclerotised, longer than wide, with 12–16 (most of which incomplete) spiral ridges ( Fig. 6D ). Three tooth-like projections situated at base of buccal capsule ( Fig. 6C ). Basal ring short and narrow, oesophageal cup poorly developed. Muscular oesophgus club-shaped, almost cylindrical in anterior half with elongated posterior bulb. Glandular oesophagus as long as muscular one, almost cylindrical along whole length, slightly widening posteriorly. Nerve ring encircling muscular oesophagus somewhat anterior to its mid-length. Position of excretory pore varying within level of muscular oesophagus posterior quarter. Intestine and rectum strait, narrow. Tail tapering in males and narrowing with six mucrons in females.

Males ( Fig. 6A ). Posterior end coiled ventrally, caudal alae relatively long, supported by papillae: 11 precloacal pedunculated papillae, two pair of adcloacal papillae (anterior and posterior to cloaca) and four pairs of postcloacal papillae ( Fig. 6F ).

Females ( Fig. 6E ). Vulva with poorly sclerotised walls, situated at mid-body level ( Fig. 6G ). Tail relatively short, narrowing, bearing six mucrons ( Fig. 6H ).

Remarks. Due to the lack of gravid specimens in our material we could not provide measurements for this species, although all morphological characters correspond to the redescription provided by Jackson and Tinsley (1995). Moravec et al. (2006) considered the genus Batrachocamallanus as junior synonym of Procamallanus . Contrary to that opinion we prefer to assign found species to the genus Batrachocamallanus due to the distant phylogenetic relationships between B. xenopodis and P. pseudolaeviconchus (14% (110 nt) in the 28S rDNA gene) and close relationships between B. xenopodis and the type species of the genus – B. slomei (4% (30 nt) in the 28S rDNA gene) (see Table 1).

Figures

Fig. 1. Camallanus sodwanaensis n. sp., line-drawings. A – anterior part of body, female, lateral view; B – buccal capsule, female, lateral view; C – anterior part of body, female, apical view; D – posterior part of body, male, ventral view; E – dorsal trident, male, lateral view; F – posterior part of body, female, lateral view; G – spicules, lateral view. Scale bars: A – 500; B–D, F–G – 100; E – 50.

Fig. 2. Camallanus sodwanaensis n. sp., photomicrographs. A – male, general view; B – anterior part of body, female, lateral view; C – buccal capsule, female, lateral view; D – optical section at level of buccal capsule valves mid-width, male, dorsal view; E – dorsal trident, male, dorsal view; F – anterior part of body, female, apical view; G - optical section at level of buccal capsule valves mid-length, male, apical view; H – right spicule, lateral view; I – posterior end of body, male, ventral view; J – part of body at vulva region, lateral view; K – posterior end of body, female, lateral view. Scale bars: A – 1 mm, B – 500, C–K – 100.

Fig. 3. Paracamallanus cyathopgharynx, photomicrographs. A – anterior part of body, male, lateral view; B – buccal capsule, male, lateral view; C – anterior part of body, male, apical view; D – female, general view; E - optical section at level of buccal capsule valves mid-length, male, dorsal view; F - part of body at vulva region, lateral view; G – posterior end of body, female, lateral view; H – posterior end of body, male, lateral view. Scale bars: A–C, E–H – 100; D – 1 mm.

Fig. 4. Procamallanus pseudolaeviconchus, photomicrographs. A – male, general view; B – anterior part of body, male, lateral view; C – anterior part of body, male, apical view; D – buccal capsule, female, lateral view; E – female, general view; F – posterior part of body, female, lateral view; G – part of body at vulva region, lateral view; H – posterior end of body, male, lateral view. Scale bars: A, E – 1 mm, B, D, F–H – 100; C – 25.

Fig. 5. Spirocamallanus daleneae, photomicrographs. A – male, general view; B – anterior part of body, male, apical view; C – female, genetal view; D – optical section at buccal capsule mid-length, male, apical view; E – optical sections at buccal capsule mid-width level, male, lateral view; F – optical section at buccal capsule 2/3 width level, male, lateral view; G – posterior end of body, male, ventral view; H – posterior end of body, female, lateral view; I – part of body at vulva region, lateral view. Scale bars: A, C – 1 mm; B, D–I – 100.

Fig. 6. Batrachocamallanus xenopodis, photomicrographs. A – male, general view; B – anterior part of body, male, apical view; C – optical section at base of buccal capsule level, male, apical view; D – buccal capsule, female, lateral view; E – female, general view; F – posterior part of body, male, lateral view; G – part of body at vulva region, lateral view; H – posterior part of body, female, lateral view. Scale bars: A, E, F–H – 100, B–D – 50.