Neofoleyellides steyni, Kuzmin & Netherlands & du Preez & Svitin, 2021
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publication ID |
https://doi.org/ 10.1016/j.ijppaw.2021.02.018 |
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persistent identifier |
https://treatment.plazi.org/id/6A14DE43-FF8D-FFAA-401E-FA6FFE9BFC98 |
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treatment provided by |
Felipe |
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scientific name |
Neofoleyellides steyni |
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Genus Neofoleyellides Netherlands, Svitin, Smit et Du Preez, 2020.
Neofoleyellides steyni n. sp.
Type-host: Amietia delalandii (Dum´eril et Bibron, 1841) ( Amphibia: Anura : Pyxicephalidae ).
Site of infection: body cavity, subcutaneous.
Type-locality: Louis Trichardt , Limpopo Province, South Africa. Coordinates: 23 ◦ 05 ′ 32.6 ′′ S, 30 ◦ 07 ′ 26.5 ′′ E GoogleMaps .
Type-specimens: Holotype ( NMB P742 View Materials ), allotype ( NMB P743 View Materials ) and 36 paratypes ( NMB P744–745 View Materials ), deposited in the Parasitic Worm Collection , National Museum , Charles Street, Bloemfontein, South Africa.
Infection parameters: Intensity 1–54 (mean 11.8); prevalence 71%; mean abundance 8.4.
Representative DNA sequences: 18S [MW599274], COI [MW598467].
Etymology: The species named after Dr Koos Steyn who provided tremendous support with sample collection.
ZooBank registration: To comply with the regulations set out in article
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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 Neofoleyellides steyni n. sp. is urn:lsid:zoobank.org:act:[228104A4-D93C-4D89-B343- FB337D8B9E5B].
Description ( Figs. 1–3 View Fig View Fig View Fig ).
General. Body thread-like, tapering on both extremities ( Fig. 1A, B, I View Fig , 2A View Fig ). Females about 2–3 times longer than males. Anterior end rounded. Body cuticle finely transversely striated. Lateral alae present in both sexes, beginning at level of mid-length of muscular oesophagus ( Fig. 3A View Fig ), almost reaching tail tip posteriorly. Alae narrow ( Fig. 3B View Fig ), triangular in transverse sections along most of body length, slightly inflated and folded in anterior and posterior regions in females ( Figs. 1I View Fig and 3A View Fig ), more prominently inflated in caudal region of males.
Oral opening oval, elongated dorso-ventrally ( Fig. 1D View Fig ). Two small lateral parastomal structures beside oral opening. One pair of large cephalic papillae with rounded tips located sub-laterally, slightly shifted to ventral side; one pair of smaller papillae, flattened and less distinct, located laterally closer to oral opening; amphidial openings pore-shaped, distinct, sub-lateral in position, slightly shifted to dorsal side ( Fig. 1D View Fig ). Buccal capsule small, cuticularised, wider anteriorly, dorso-ventrally elongated in apical view ( Fig. 1C, E, F View Fig ).
Muscular oesophagus comparatively short and narrow, rounded on anterior end ( Fig. 1C View Fig ), widening posteriorly, with slight constriction in posterior part, just anterior to junction with glandular oesophagus ( Fig. 1A and B View Fig ). Lumen of muscular oesophagus tri-radiate ( Fig. 1G View Fig ). Border between glandular and muscular oesophagus indistinct. Glandular oesophagus wider and approximately 3–4 times longer than muscular oesophagus, its anterior half gradually widening posteriorly, posterior half narrowing ( Fig. 1A and B View Fig ). Nerve ring surrounding constriction in posterior part of muscular oesophagus.
Males (30 specimens) Table 2. Body 17–26 [19] mm long, 130–209 [142], 217–316 [217] and 203–312 [235] wide at nerve ring, oesophageal-intestinal junction and mid-body, respectively. Posterior part of body corkscrew-shaped. Lateral alae beginning at 50–166 [85] from anterior end. Buccal capsule 4–7 [4] long, 5–10 [7] wide. Muscular portion of oesophagus 219–507 [270] long, 26–48 [27], 26–49 [35] and 22–54 [52] wide at anterior, mid-length and posterior level, respectively. Glandular portion of oesophagus 1032–1813 [1125] long, 29–75 [67], 80–155 [99] and 48–88 [60] wide at anterior, mid-length and posterior level, respectively. Total length of oesophagus 1318–2217 [1395], or 6.3–11.9 [7.5] % of body length. Nerve ring located at 167–351 [245] from anterior end; this distance corresponding to 10.7–19.4 [17.6] % of total oesophagus length. Anterior part of testis narrow, forming loops and bends in region of glandular oesophagus ( Fig. 1B View Fig ). Posterior to oesophago-intestinal junction, testis wide and straight. Ejaculatory duct narrower than testis, with funnel-shaped proximal part.
In caudal region, lateral alae differing in shape and size: left ala larger and more inflated ( Fig. 3C View Fig ). Area rugosa present on ventral side of body in posterior part, consisting of rounded bosses arranged in somewhat irregular transverse rows ( Fig. 3D View Fig ). Bosses not reaching cloacal region posteriorly ( Fig. 2A View Fig ). Single cuticular elevated plaque present anterior to cloacal aperture, often slightly shifted to right side. Tail short, conical, with rounded tip, 47–76 [55] long. In most specimens, five pairs of ventro-lateral genital papillae present: four pairs of large and rounded papillae (one pre-cloacal and three post-cloacal) and one pair of smaller sessile papillae located closer to posterior end ( Fig. 2D View Fig ). Papillae on left and right side not always symmetrical in position ( Fig. 2E View Fig ). In some specimens, number of papillae, on at least one side,
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reduced to four ( Fig. 2F and G View Fig ), or increased to six on one or both sides ( Fig. 2H and I View Fig ) due to presence of additional pre- and/or post-cloacal papillae. Phasmids small, subterminal. Spicules unequal and dissimilar ( Fig. 2A View Fig ). Right spicule short and robust, 72–127 [105] long, slightly curved ventrally, with funnel-shaped proximal end separated from spicule body by apparent constriction ( Fig. 2B View Fig ). Distal end of spicule rounded. Left spicule longer and thinner, 196–314 [287] long, curved ventrally, alate, with rough surface of body and smooth blade; ala almost reaching distal end of spicule ( Fig. 2C View Fig ). Proximal part of spicule not distinctly separated from spicule body.
Females (30 specimens) Table 3. Body 34–68 [52] mm long, 165–313 [185], 311–482 [426] and 306–552 [433] wide at nerve ring, oesophageal-intestinal junction and mid-body length, respectively. Lateral alae beginning at 70–221 [112] from anterior end. Buccal capsule 5–7 [5] long, 4–9 [8] wide. Muscular portion of oesophagus 267–483 [326] long, 31–49 [46], 28–51 [44] and 33–61 [41] wide at anterior, mid-length and posterior level, respectively. Glandular portion of oesophagus 1413–2421 [1817] long, 41–93 [59], 98–205 [115] and 55–164 [75] wide at anterior, mid-length and posterior level, respectively. Total length of oesophagus 1784–2733 [2143], or 3.1–5.7 [4.2] % of body length. Nerve ring situated at 231–373 [261] from anterior end; this distance corresponding to 10.4–20.0 [12.2] % of total oesophagus length.
Vulva not salient, transverse, located at level of anterior part of glandular oesophagus ( Fig. 1A View Fig ), at 508–1090 [752] from anterior end of body, this distance corresponding to 1.0–2.1 [1.5] % of body length and 24.3–56.0 [35.1] % of total oesophagus length. Vagina short, globular. Vagina vera infundibuliform, lined with thin cuticle. Vagina uterina with thick cuticular lining, curved, connected to thick-walled ovejector. Ovejector directed posteriorly, then forming bends and loops along and around oesophagus ( Fig. 1A View Fig ). Uteri and ovaries forming numerous loops and filling whole body of female, not reaching caudal region. Intestine narrow in posterior part. Rectum thin, short, cuticularised. Tail gradually tapering, with rounded tip ( Figs. 1I View Fig ), 101–277 [153] long.
Microfilariae (30 specimens). Sheath often visible around posterior part of microfilaria. Body short, gradually tapering in posterior half ( Figs. 1H View Fig and 7A View Fig ) and 70–112 long and 3–7 wide. Anterior end rounded, posterior end pointed. Cephalic hook minute. Crown of about 20 short cuticular ridges present on anterior extremity. Densely arranged chromatin, staining dark purple with Giemsa-stain ( Fig. 7A View Fig ).
Remarks. The new species was assigned to the genus Neofoleyellides based on the morphology of apical extremity: oral opening small and surrounded by two lateral parastomal structures, 2 minute pore-like amphids, two small internal lateral papillae and two enlarged external sublateral papillae; in males: presence of caudal alae, wide plaque, and unequal spicules; in females: vulva situated at level of oesophagus; microfilariae with sheath ( Netherlands et al., 2020).
Neofoleyellides steyni n. sp. can be distinguished from N. boerewors by asymmetrical caudal alae in males and the asymmetry in number and position of genital papillae, contrary to narrow and similar alae and four papillae on each side in N. boerewors . Two species have different position of the vulva in females: at level of anterior end of the glandular portion of the oesophagus in N. steyni n. sp. (24–56% of total oesophagus length)
vs closer to posterior end of the glandular portion of the oesophagus (68–103% of total oesophagus length) in N. boerewors . The shape of the glandular portion of the oesophagus in both sexes is also different in the two species: glandular oesophagus of N. steyni n. sp. has maximum width at level of the border between the first and second thirds, and narrowing posterior third while glandular oesophagus in N. boerewors is evenly
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widened, with maximum width at posterior end.
Neofoleyellides steyni n. sp. clearly differs from N. martini n. sp. in morphology of caudal alae in males, position of the vulva and morphology of microfilariae (see below).
Family Onchocercidae Leiper, 1911
Subfamily Waltonellinae Bain et Prod’ Hon, 1974.
Genus Neofoleyellides Netherlands, Svitin, Smit et Du Preez, 2020.
Neofoleyellides martini n. sp.
Type-host: Leptopelis natalensis (Smith, 1849) ( Amphibia: Anura : Arthroleptidae ).
Site of infection: body cavity, subcutaneous, pericardium.
Type-locality: Alfred Park , Pinetown, KwaZulu-Natal Province, South Africa. Coordinates: 29 ◦ 47 ′ 30.4 ′′ S 30 ◦ 52 ′ 06.0 ′′ E GoogleMaps .
Type-specimens: Holotype ( NMB P739 View Materials ), allotype ( NMB P740 View Materials ) and 20 paratypes ( NMB P741 View Materials ), deposited in the Parasitic Worm Collection,
National Museum, Charles Street, Bloemfontein, South Africa.
Infection parameters: All of eight dissected amphibians were infected with 3–54 (average 15) nematodes.
Representative DNA sequences: 18S [MW599275], COI [MW774895].
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 Neofoleyellides martini n. sp. is urn:lsid:zoobank.org:act:[30C787D3-9B0A-4B47-A32B- 1AADE504C855].
Etymology: The species is named after Henry Martin (close family friend to author ECN), who provided support and accommodation for the team during several sampling trips to the area and assisted in the initial discovery of this species.
Description ( Figs. 4–6 View Fig View Fig View Fig ).
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General. Body thread-like, tapering on both extremities ( Fig. 4A, B, G, I View Fig ). Females about 2–3 times longer than males. Anterior end rounded. Body cuticle finely transversely striated. Lateral alae present in both sexes, beginning at level of mid-length of muscular oesophagus, almost reaching tail tip posteriorly. Alae narrow, triangular in transverse sections along most of body length, slightly inflated and folded in anterior and posterior regions.
Oral opening oval, elongated dorso-ventrally ( Fig. 4D View Fig ). Two small lateral parastomal structures beside oral opening ( Fig. 4D and E View Fig ). One pair of large cephalic papillae with rounded tips located sub-laterally, slightly shifted to ventral side; one pair of smaller papillae, flattened and less distinct, located laterally closer to oral opening; amphidial openings pore-shaped, distinct, sub-lateral in position, slightly shifted to dorsal side ( Fig. 4 C–F View Fig ). Buccal capsule inconspicuous.
Muscular oesophagus short and narrow, rounded on anterior end, widening posteriorly ( Fig. 4A and B View Fig ). Lumen of muscular oesophagus tri-radiate ( Fig. 4F View Fig ). Border between glandular and muscular oesophagus indistinct. Glandular oesophagus wider and approximately 5–6 times longer than muscular oesophagus, its anterior half gradually widening posteriorly, posterior third narrowing ( Fig. 4A and B View Fig ). Nerve ring at mid-
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length of muscular oesophagus or slightly posterior to it.
Males (30 specimens) Table 2. Body 14–19 [16] mm long, 113–146 [116], 202–263 [256] and 214–292 [235] wide at nerve ring, oesophageal-intestinal junction and mid-body, respectively. Posterior part of body corkscrew-shaped. Lateral alae beginning at 42–175 [97] from anterior end. Muscular portion of oesophagus 203–308 [225] long, 24–40 [36], 29–51 [35] and 27–41 [29] wide at anterior, mid-length and posterior level, respectively. Glandular portion of oesophagus 972–1607 [1201] long, 42–79 [45], 74–125 [89] and 51–97 [80] wide at anterior, mid-length and posterior level, respectively. Total length of oesophagus 1237–1862 [1426], or 7.5–11.7 [8.7] % of body length. Nerve ring located at 186–258 [191] from anterior end, this distance corresponding to 11.3–18.7 [13.4] % of total oesophagus length. Anterior part of testis narrow, forming loops and bends in region of glandular oesophagus ( Fig. 4B View Fig ). Posterior to oesophago-intestinal junction, testis wide and straight. Ejaculatory duct narrower than testis, with funnel-shaped proximal part.
In caudal area, lateral alae equal, slightly inflated, semi-spherical on transverse section ( Fig. 6A View Fig ). Area rugosa present on ventral side of body in posterior part, consisting of rounded bosses arranged in somewhat irregular transverse rows ( Fig. 6B View Fig ). Bosses not reaching cloacal region posteriorly. Single cuticular elevated plaque present anterior to cloacal aperture. Tail short, conical, with rounded tip, 41–76 [55] long. Number and position of genital papillae somewhat variable. In most males (16 out of 30), 12 ventrolateral papillae arranged in two groups: four pairs of large, mammiform papillae in anterior group, of them one pair distinctly precloacal, one pair pre- or ad-cloacal, and two pairs post-cloacal, and two pairs of small papillae in posterior group, closer to posterior extremity ( Figs. 4G View Fig and 5A View Fig ). In some specimens (12 out of 30), one papilla in anterior-most pair absent on either side ( Fig. 5B View Fig ). In few specimens, absence of anterior-most papilla accompanied with absence of both posterior papillae on same side, while remaining posterior papillae enlarged and located closer to anterior group ( Fig. 5C and D View Fig ). Phasmids pore-like, subterminal ( Figs. 4G View Fig , 5A–D View Fig ). Spicules unequal and dissimilar ( Fig. 4J View Fig ). Right spicule short and robust, 113–159 [140] long, slightly curved ventrally, with funnel-shaped proximal end separated from spicule body by apparent constriction. Distal end of spicule rounded. Left spicule longer and thinner, 275–399 [358] long, curved ventrally, with rough surface of body and smooth blade with sharpened tip. Proximal part of spicule not distinctly separated from spicule body.
Females (30 specimens) Table 3. Body 17–37 [37] mm long, 146–197 [197], 266–468 [468] and 289–551 [447] wide at nerve ring, oesophageal-intestinal junction and mid-body length, respectively. Lateral alae beginning at 65–199 [95] from anterior end. Muscular portion of oesophagus 230–409 [409] long, 24–44 [43], 27–42 [38] and 27–42 [33] wide at anterior, mid-length and posterior level, respectively. Glandular portion of oesophagus 1319–2076 [2076] long, 39–77 [66], 75–142 [110] and 50–113 [82] wide at anterior, mid-length and posterior level, respectively. Total length of oesophagus 1670–2485 [2485], or 3.1–5.7 [4.2] % of body length. Nerve ring situated at 231–373 [261] from anterior end, this distance corresponding to 5.0–10.4 [6.7] % of total oesophagus length.
Vulva not salient, transverse, located at level of oesophagus mid-length ( Fig. 4A View Fig ), at 629–1340 [12,251] from anterior end of body, this distance corresponding to 2.6–4.4 [3.4] % of body length and 36.5–63.5 [50.3] % of total oesophagus length. Vagina short, globular. Vagina vera infundibuliform, lined with thin cuticle. Vagina uterina with thick cuticular lining, curved, connected to thick-walled ovejector. Ovejector directed posteriorly, then forming bends and loops along and around oesophagus ( Fig. 4A View Fig ). Uteri and ovaries forming numerous loops and filling whole body of female, not reaching caudal region. Intestine narrow in posterior part. Rectum thin, short, cuticularised. Tail gradually tapering, with rounded tip ( Figs. 4I View Fig ), 51–172 [92] long.
Microfilariae (30 specimens). Sheath present as loose envelope around whole body of larva. Body short and stout, rounded on anterior end, tapering in posterior half, 51–79 long and 5–9 wide ( Figs. 4H View Fig and 7B View Fig ). Densely arranged chromatin, staining dark purple to pink with Giemsa-stain, slight dilatation apparent just posterior to body midregion due to presence of large translucent G1 cell ( Figs. 4H View Fig and 7B View Fig ). Cephalic hook and crown of 10–15 short cuticular ridges on apical surface.
Remarks. The new species was assigned to the genus Neofoleyellides based on morphology of apical extremity: oral opening small and surrounded by two lateral parastomal structures, 2 minute pore-like amphids, two small internal lateral papillae and two enlarged external sublateral papillae; in males: presence of caudal alae, wide plaque, and unequal spicules; in females: vulva situated at level of oesophagus; microfilariae with sheath ( Netherlands et al., 2020).
Neofoleyellides martini n. sp. can be differentiated from N. boerewors by arrangement of caudal papillae in males: often asymmetrical, with first pair pre-cloacal in N. martini n. sp. contrary to symmetrical with first pair ad-cloacal in N. boerewors . Females of N. martini n. sp. have vulva located near oesophagus mid-length (37–64% of oesophagus length) in contrast to females of N. boerewors having vulva at level close to oesophagus posterior end (68–103%).
Neofoleyellides martini n. sp. differs from N. steyni n. sp. in the shape of male caudal alae, relative position of vulva in females and morphology of microfilariae. In males of N. martini sp. n., the caudal alae are narrow and symmetrical, while in N. steyni n. sp., the left ala is prominently larger than the right one. In females of N. martini n. sp., the vulva is usually located more posteriorly (37–64% of total oesophagus length; 3–4% of body length) than in females of N. steyni n. sp. (24–56%
of total oesophagus length; 1–2% of body length), though this character is variable and the values are overlapping in both species. The
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microfilariae in N. martin n. sp. are shorter (51–79 long), with large G1
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