Hemicycliophora undetermined

Subbotin, Sergei A., Chitambar, John J., Chizhov, Vlamidir N., Stanley, Jason D., Inserra, Renato N., Doucet, Marcelo E., Clure, Michael M, Ye, Weimin, Yeates, George W., Mollov, Dimitre S., Cantalapiedra-Navarrete, Carolina, Vovlas, Nicola, Berg, Esther Van Den & Castillo, Pablo, 2014, Molecular phylogeny, diagnostics, and diversity of plant-parasitic nematodes of the genus Hemicycliophora (Nematoda: Hemicycliophoridae), Zoological Journal of the Linnean Society 171 (3), pp. 475-506 : 482-491

HEMICYCLIOPHORA SPECIES

Brief morphological descriptions with illustrations ( Figs 1–5, S1–S 17) and morphometric values ( Tables S1– S 8) are given for the populations of the earlier mentioned 20 Hemicycliophora species. All topotype populations of H. floridensis , H. halophila , H. hellenica , H. iberica , and H. italiae were compared with paratypes and were coincident with the original descriptions (viz. Chitwood & Birchfield, 1957; Yeates, 1967; Vovlas, 2000; Castillo et al., 1989; Brzeski & Ivanova, 1978; respectively) in their morphology and morphometrics ( Figs 2A, F, K , 4, S4 , S 5 ; Table S2).

HEMICYCLIOPHORA CALIFORNICA BRZESKI, 1974

( FIG. 1F, L, R, TABLES 1, S1)

Hemicycliophora californica was first found in association with apricot roots in southern California. Morphological analysis of a population from Yolo County, California, showed that females were characterized by a straight or ventrally arcuate body, lateral fields marked by breaks and anastomoses of transverse striae, forming an occasional discontinuous, single longitudinal line over short distances in mid- and prevulval body regions, annuli outside lateral fields smooth or inconsistently marked with fine lines or ridges, lip region round to hemispherical with three annuli, labial disc slightly protruding and rounded in lateral view, sometimes low and flat, vulval lips modified (slightly elongated), postvulval body constricted immediately behind vulva then cylindrical to anus and tail cylindrical then tapering abruptly to a uniformly conical posterior third region, occasionally with a greater dorsal curvature, with a short conical to slightly attenuated conical spike and narrowly rounded terminus. Tail terminus distinctly annulated.

Except for the infrequent presence of fine lines or ridges and occasional irregular lines on annuli outside the lateral fields, and three lip annuli (vs. two annuli), the morphology and morphometrics of the Californian population of H. californica studied here agreed well with the original and subsequent descriptions by Brzeski (1974) and Costa-Manso (1996), respectively. This species has been found only in California and Idaho, USA.

Distinguishing characters include a conoid tail with a narrowly rounded to acute terminus, and presence of males. Hemicycliophora californica is differentiat- ed from the morphologically similar species H. raskii and Hemicycliophora montana Eroshenko, 1980 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA CONIDA THORNE, 1955

( FIGS 1A, G, M, S1 ; TABLES 1, S1)

Hemicycliophora conida was described by Thorne from a population collected from sugarbeet near Ballyculane, Ireland. Two populations of this species found in Spain and Washington, USA, are described in this study. Females were characterized by a straight or ventrally arcuate body, lateral fields marked by breaks and anastomoses with three longitudinal lines starting as a distinct central line from just posterior to pharyngeal base to near the midbody region where two faint lines appear adjacent on either side forming two rows of blocks with the transverse striae, lip region with rounded anterior margins, two distinct annuli, three annuli suggested with first annulus less distinct and narrower than remaining two distinct lip annuli, vulval lips modified, vulval sheath about one to two annuli long and annuli at tail tip smaller than at anterior tail. No males were found.

Morphology and morphometrics of the Spanish and Washington populations were coincident with those provided for this species ( Thorne, 1955; Loof, 1968), except for minor intraspecific morphometric differences (e.g. body length 683–912 vs. 660–990 μm; stylet 70–89 vs. 73–96 μm). Hemicycliophora conida has been widely reported in European countries ( Loof, 1968; Brzeski, 1974, 1998; Peña-Santiago et al., 2004) and Iran ( Loof, 1984). This is the first report of H. conida from the Americas. The specimens recently identified as H. conida by Zeng et al. (2012) from grasses in North and South Carolina do not fully fit the original and other H. conida descriptions [e.g. tail length 100.5– 128.2 vs. 60–93 μm; Rex (number of annuli between anterior end of body and excretory pore) 49–67 vs. 32–52 and others] and probably belong to H. wyei or other species.

Distinguishing characters of this species include lateral fields marked by anastomoses bordered on both sides by rows of ornamentations, R 175–278, stylet 69–101 μm, a short conoid tail ending in a narrowly rounded to acute terminus, and presence of males. Hemicycliophora conida is differentiated from the morphologically similar species Hemicycliophora iwia Brzeski, 1974 and Hemicycliophora ovata Colbran, 1962 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA EPICHAROIDES LOOF, 1968 ( FIG. S2 , TABLES 1, S1)

Hemicycliophora epicharoides was described by Loof from sandy soil in the Netherlands. The populations of this species described in this study were collected from sandy soils in Italy and Spain. Females of these popu- lations were characterized by coarse body annuli (5– 7 μm wide), the number of body annuli fewer than 190, a rounded lip region (18.5–22 μm wide), lateral fields marked by breaks in the annuli, vulval lips modified, slightly elongate, and tail cylindroid, then becoming bluntly triangular distally, without sharp demarcation between the two parts. Morphologically and morphometrically the Italian and Spanish populations are similar to the original description ( Loof, 1968), except for minor differences including a looser cuticular sheath, smaller body length/maximum body width (17.1–22.2 vs. 21–27) value, and a slightly smaller tail length/ body width at anus value (1.5–2.1 vs. 2.1–2.7).

This species was found in several European countries including France, Italy, the Netherlands, Poland, and Spain ( Loof, 1968; Germani & Luc, 1973; Brzeski 1974; Vovlas & Inserra, 1980; Peña-Santiago et al., 2004), Korea ( Choi & Geraert, 1995) and South Africa ( Van den Berg & Tiedt, 2001).

Distinguishing characters of this species include lip annuli not separated from the rest of the body, stylet knobs with large cavity, lateral fields with two lines, a conoid tail with a rounded to acute terminus, and presence of males. Hemicycliophora epicharoides is differentiated from the morphologically similar species Hemicycliophora demani Edward & Rai, 1971 , and Hemicycliophora koreana Choi & Geraert, 1971 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA GRACILIS THORNE, 1955

( FIGS 3E, S3 ; TABLES 1, S 2)

Hemicycliophora gracilis was described by Thorne from specimens collected in five states in the USA. The populations described in this study are from California and one from Minnesota. Females of these populations were characterized by a ventrally arcuate body, cuticular sheath loosely fitting, often extending over anterior end, lateral fields marked by breaks and anastomoses in between two longitudinal lines, annuli smooth outside lateral field, sometimes marked with few short, irregular longitudinal lines that mark midbody region near lateral fields, lip region rectangular to slightly hemispherical, three lip annuli with first annulus narrower than second and third lip annuli and often difficult to discern, labial disc slightly elevated, vulval lips modified, about two annuli long, tail anteriorly cylindrical then tapering uniformly to an attenuated conical posterior third with almost cylindrical attenuation with a rounded terminus. Annulation more or less distinct on terminus. One male was found ( Fig. S3 ). This is the first report of males for this species .

The Californian population of H. gracilis closely resembles other populations reported elsewhere in the USA ( Thorne, 1955; Brzeski, 1974), but differs from them by a slightly shorter female body (1044–1212 vs. 1230–1700 μm), shorter stylet (91–116 vs. 111– 132 μm), smaller R (322–380 vs. 341–395), and three vs. two lip annuli.

Distinguishing characters of this species include a long body (1.2–1.5 mm) and stylet (111–132 μm), posterior part of tail distinctly offset with a spicate terminus, and presence of males. Hemicycliophora gracilis is differentiated from the morphologically similar species Hemicycliophora litorea Van den Berg, 1988 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA IBERICA CASTILLO , GÓMEZ- BARCINA & LOOF, 1989

( FIG. 4 ; TABLES 1, S 2)

Two new Spanish populations of H. iberica from Hinojos , Huelva Province , and Santa Elena , Jaén Province , were detected and found to be similar in morphology and morphometrics to the original description ( Castillo et al., 1989), except for minor differences such as a shorter stylet (80–91 vs. 79.94 μm) and a smaller Rst (number of annuli between anterior end of body and stylet base) value (15–27 vs. 18–31). As the species was only known from the type locality, the new records of the species extend its distribution within southern Spain .

Distinguishing characters of this species include two incisures, truncated lip region with three annuli, a long stylet, absence of males, tail elongate-triangular with distal part offset ( Castillo et al., 1989).

HEMICYCLIOPHORA LUTOSOIDES LOOF, 1984 ( FIG. S6; TABLES 1, S 3)

Hemicycliophora lutosoides was described by Loof (1984) from populations collected in Iran and Iraq. Two female populations of H. lutosoides from San Pablo de Buceite, Cádiz, and Los Palacios y Villafranca, Seville Province in Spain are described here. Females of these populations were characterized by a truncate-rounded lip region with a barely protruding labial disc, body annuli more than 300, lateral fields marked by breaks, vulval lips modified, and an elongate tail tapering uniformly to an acute terminus. Morphology and morphometrics of these populations were similar to the original description ( Loof, 1984), except for minor intraspecific differences including a smaller number of body annuli (303–364 vs. 297–376) and a smaller Rex (54–65 vs. 57–70). The present record of H. lutosoides is the second one from Spain, after Bello (1979), and the third after its original description. An additional population with similar morphometrics was collected in Hinojos, Huelva Province, Spain, associated with wild olive.

Costa-Manso (1998) synonymized this species with Hemicycliophora lutosa Loof & Heyns, 1969 , from South Africa considering described differences in body and tail annuli as intraspecific variations. However, our molecular data do not support this synonymy and indicate that H. lutosoides is a valid species.

Distinguishing characters of this species include a truncated and not elevated labial disc, lateral fields marked by breaks and anastomoses, slightly elongat- ed vulval lips, a conical postvulval region, a uniformly tapering elongate conoid tail with an acute terminus, and absence of males. Hemicycliophora lutosoides is differentiated from the morphologically similar species H. lutosa ( Chitambar & Subbotin, 2014) .

HEMICYCLIOPHORA OBTUSA THORNE, 1955

( FIG. 5 ; TABLES 1, S 3)

Hemicycliophora obtusa was described by Thorne from a population found in Utah, USA. The population described in this study was found in the rhizosphere of Scotch pine growing in sandy soil in Spain. Females of this population were characterized by a cuticular sheath closely adpressed to the cuticle, lip region continuous with body and bearing two annuli, labial disc undistinguishable, lateral fields with anastomoses or breaks in striae, vulval lips modified, slightly modified and tail typically hemispheroid with rounded terminus and usually smooth tip (although annulation was observed in some specimens). Spermatheca almost hemispherical (20–22 μm wide), filled with rounded sperm cells, 1.5–2.0 μm in diameter. Males very rare, only a pre-adult stage was detected without a stylet, a rounded tail tip, a short bursa, and semicircular spicules. The morphology and morphometrics of the Spanish population agree with the original description and redescription ( Thorne, 1955; Brzeski, 1974), except for minor intraspecific differences: R (190–219 vs. 219– 268), VL/VB (distance between vulva and posterior end of body divided by body width at vulva) (2.9–4.2 vs. 3.0–4.4), V [(distance from head end to vulva/ body length)*100] (82–86 vs. 86–90%), and a slightly shorter stylet (62–88 vs. 78–98 μm). Hemicycliophora obtusa is known only from the type population from Utah, USA. The present record of H. obtusa is the first from Europe and the second world record after the original description. An additional population with similar morphometrics was collected in Sanlúcar de Barrameda, Cádiz Province, Spain, associated with wild olive.

Distinguishing characters of this species include the absence of longitudinal lines outside lateral fields, lateral fields without lines, a hemispherical tail terminus, and absence of males. Hemicycliophora obtusa is differentiated from the morphologically similar species H. arenaria ( Chitambar & Subbotin, 2014) .

HEMICYCLIOPHORA PORANGA MONTEIRO & LORDELLO, 1978

( FIGS 2B, G, L , 3B, I, S 7 ; TABLES 1, S4)

Hemicycliophora poranga was described by Monteiro and Lordello from a population in a cabbage field in Brazil. The populations examined in this study were collected in Italy, Spain, and California, USA. Females of these populations were characterized by a rounded to hemispherical lip region in lateral view with an elevated labial disc. Lateral fields marked by breaks and anastomoses, which, in some populations, are between two more or less distinct longitudinal lines starting near base of pharynx and extending to the posterior part of the body. Vulval lips modified, about one to two annuli long, posteriorly directed with short vulval sleeve. Tail tapering uniformly to a narrow, attenuated conical posterior part with a finely rounded terminus. Annulation on tail almost distinct up to terminus. Morphology and morphometrics of Italian and Spanish populations are generally similar to the original description ( Monteiro & Lordello, 1978). The present records of H. poranga are the first from Italy and Spain, and constitute the first records in Europe. The species has been reported from Argentina ( Doucet, 1982; Chaves, 1983), Iran ( Jamali et al., 2004), Venezuela ( Crozzoli & Lamberti, 2006), and California, USA ( Chitambar, 1994).

Distinguishing characters of this species include an elevated labial disc and elongate conoid tail uniformly tapering to a narrowly rounded to acute terminus, and presence of males. Hemicycliophora poranga is differentiated from the morphologically similar species Hemicycliophora ornamenta Bajaj, 1998 , H. ripa Van den Berg, 1981 , Hemicycliophora subaolica Jairajpuri & Baqri, 1973 , and Hemicycliophora micoletzkyi Goffart, 1951 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA RASKII BRZESKI, 1974

( FIG. 1D, J, P; TABLES 1, S3)

Hemicycliophora raskii was described by Brzeski from a population collected from the rhizosphere of almond trees in Oakley , Contra Costa County, California, USA. This species is known only from California. Another population was found in Sacramento County , California, and described in this study. Females of this population were characterized by a slightly ventrally arcuate body, cuticular sheath closely or loosely fitting, lateral fields marked by anastomoses and continuous transverse striae – a short, irregular longitudinal line was observed in one specimen, lip region rounded to hemispherical with three annuli usually with first lip annuli faintly visible labial disc not protruding, vulval lips modified, about one annulus long posteriorly direct- ed, tail cylindrical then abruptly conoid to wedgeshaped in posterior third tapering uniformly to a narrow, slightly attenuated spike with a finely rounded terminus; variable forms having a more conoid posteri- or region with a wider conical spike or with a rounded terminus .

The population closely resembles the original description of H. raskii ( Brzeski, 1974) in lip region shape and number of annuli, markings of the lateral fields and annuli, shape of vulval lips and tail. The presence of distinct irregular annular lines outside and near the lateral fields have not been reported earlier for this species, and the studied population differed from the original description by a slightly shorter stylet (77– 85 vs. 86–94 μm), slightly larger R (194–234 vs. 166– 201), and RV-anterior end (number of body annuli from anterior end to vulva) (161–194 vs. 134–163).

Distinguishing characters of this species include the presence of longitudinal lines or scratches outside lateral fields, very small stylet knobs with very small or no cavity, a conical to wedge-shaped tail with a finely rounded terminus, and absence of males. Hemicycliophora raskii is differentiated from the morphologically similar species H. californica ( Chitambar & Subbotin, 2014) .

HEMICYCLIOPHORA RIPA VAN DEN BERG, 1981 ( FIG. S9 ; TABLES 1, S 3)

Hemicycliophora ripa was described by Van den Berg from a population found in South Africa. A popula- tion of this species was also found in Spain and described in this study. Females of the Spanish population were characterized by a loosely fitting cuticular sheath and typical cuticular ornamentation, appearing as blocks inside annuli, a rounded lip region with two annuli and an elevated labial disc, 7.7 ± 0.6 (7.0–8.0) μm wide, body annuli more than 240, lateral fields marked by anastomoses or breaks, vulval lips modified, and an elongate tail tapering uniformly to an acute terminus. Morphology and morphometrics of the Spanish population are coincident with the original description ( Van den Berg, 1981) and those of Argentinean populations ( Doucet, 1982; Chaves, 1983) except for minor intraspecific differences including a slightly wider lip region (20–23 vs. 16.2–21.1 μm), a slightly smaller number of body annuli (229–248 vs. 246–288, 241– 316), a slightly longer stylet (94–110 vs. 81.5–98.6, 65–97 μm), a slightly larger VA%T value (54.2–86.3 vs. 25.1–70.7, 35–65), and a slightly shorter tail (78–103 vs. 84–172, 60–113 μm). The present record of H. ripa is the first from Spain, and constitutes the first record in Europe.

Distinguishing characters of this species include a loosely fitting cuticular sheath, lateral fields marked only by anastomoses or breaks in the striae occasionally forming a faint line, and the absence of males. Hemicycliophora ripa is distinguished from the morphologically similar species H. subaolica , Hemicycliophora popaensis Van den Berg & Tiedt, 2005 , and Hemicycliophora catarinensis Costa-Manso, 1996 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA SIMILIS THORNE 1955 ( FIG. S10 ; TABLES 1, S 3)

Hemicycliophora similis was described by Thorne from agricultural and noncultivated fields in the Pacific North West, USA. A population of this species was found in strawberry rhizosphere soil in Cartaya, Huelva Province, Spain and described in this study. This Spanish population was characterized by a rounded lip region with two or three annuli and a barely protruding labial disc (6.0–6.5 μm wide), body annuli around 300, lateral fields marked by breaks or anastomoses of body annuli without longitudinal lines, stylet knobs rounded, without distinct cavity, hemizonid usually undistinguishable, occupying two annuli; vulval lips not modified (not elongated, rounded) and tail with a conoid-rounded end, clearly offset with a terminal somewhat elongate spike (39–52 μm long). Morphology and morphometrics of this population were similar to those of the original description and redescription ( Thorne, 1955; Brzeski, 1974), except for minor intraspecific differences including a slightly longer body (1022–1578 vs. 1110–1200 μm) and stylet (91–108 vs. 88–96 μm), and larger VL/VB ratio (4.8–7.6 vs. 4.1–5.2). The present record of H. similis is the first for Europe and Spain, and the second after the original description from Utah, USA ( Thorne, 1955). Arias, López-Pedregal & Jiménez-Millán (1963) and Jiménez-Millán et al. (1965) recorded this species from southern Spain in Seville Province, but according to Bello (1979) this material belongs to H. thornei .

Distinguishing characters of this species include a rounded labial disc, vulval lips modified and with an elongated posterior lip, an elongate conoid tail tapering to a distal, offset and narrower elongated portion with a narrowly rounded terminus, and the absence of males. Hemicycliophora similis is differentiated from morphologically similar species, Hemicycliophora filicauda Doucet, 1982 , H. thornei , H. vaccinii , and Hemicycliophora zuckermani Brzeski, 1963 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA THIENEMANNI (SCHNEIDER, 1925) LOOS, 1948 (= LOOFIA THIENEMANNI (SCHNEIDER, 1925) SIDDIQI, 1980 ) (FIG. S11; TABLES 1, S5)

Hemicycliophora thienemanni was described by Schneider from a population found in Germany. Populations of this species examined in our study were found in Italy (southern), Spain (northern and southern), and Russia. These populations were characterized by a slender body, a conoid-rounded lip region with indistinct lips, lateral fields marked by breaks or anastomoses, vulval lips unmodified and tail long (88– 128 μm) with posterior end distinctly offset, annulations on posterior part of tail usually distinct but annuli smaller than other tail annuli in females. Morphology and morphometrics of the three studied populations were quite similar to each other, showing few intraspecific differences in body annuli and width of lip region, and fitted with data for several populations from the Netherlands and Poland, e.g. in stylet length (86–105 vs. 85–99, 77–106 μm, respectively) and V (80–85 vs. 81–84, 78–85%, respectively) ( Loof, 1968; Brzeski, 1974). This is probably the most widespread species of the genus, and has been reported in several European countries including Belgium, England, France, Germany, Greece, Italy, Spain, and Switzerland ( Loof, 1968; Brzeski, 1974; Boag & Orton Williams, 1976; Koliopanos & Vovlas, 1977; Peña-Santiago et al., 2004; Subbotin et al., 2005), as well as in Oregon, USA ( Hafez et al., 1992), Argentina ( Chaves, 1983), Brazil ( Rashid, Geraert & Sharma, 1987), Martinique ( Van den Berg & Cadet, 1992), Egypt ( Ibrahim, Mokbel & Handoo, 2010), and South Africa ( Van den Berg & Tiedt, 2001).

Distinguishing characters of this species include lateral fields with breaks or anastomoses, an offset tail end, and the absence of males. Hemicycliophora thienemanni is differentiated from the morphologically similar species H. thornei , H. poranga , Hemicycliophora indica Siddiqi, 1961 , and Hemicycliophora postamphidia Rahaman, Ahmad & Jairajpuri, 1966 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA THORNEI GOODEY, 1953 (FIG. S12; TABLES 1, S5)

Hemicycliophora thornei was described by Goodey from a population found in the Netherlands. The population examined in our study was found in La Rambla, Córdoba Province, Spain. This Spanish population was characterized by a conoid-rounded, slightly elevated lip region, body annuli without longitudinal lines or scratches outside lateral fields, lateral fields marked only with breaks and irregularities in the transverse striae, vulval lips modified, tail conical, end tapering and distinctly offset (Fig. S12). This species was previously reported from southern Spain by Arias et al. (1963), but no information of taxonomic interest was given about the Iberian populations. Morphology and morphometrics of the Spanish population were similar to the original description and redescriptions of specimens from the Netherlands ( Loof, 1968; Brzeski, 1974), except for minor intraspecific differences including a slightly longer stylet (94–114 vs. 89–103 μm) and slightly smaller VL/VB ratio (4.0–4.3 vs. 4.4–4.7). The species has also been reported in Germany, Poland, Spain, and Hungary ( Loof, 1968; Brzeski, 1974; Peña-Santiago et al., 2004; Andrássy, 2007).

Distinguishing characters of this species include a conoid lip region, lateral lips higher than submedian ones, a subcylindroid elongate conoid tail distally spicate and distinctly offset with a finely rounded terminus, and the presence of males. Hemicycliophora thornei is differentiated from the morphologically similar species H. thienemanni and Hemicycliophora monticola Mehta, Raski &Valenzuela, 1983 ( Chitambar & Subbotin, 2014).

HEMICYCLIOPHORA VACCINII REED & JENKINS, 1963 (= LOOFIA VACCINII ( REED & JENKINS, 1963) SIDDIQI, 1980) (FIG. S13; TABLES 1, S5)

Hemicycliophora vaccinii was described by Reed and Jenkins from a population associated with cranberry in Massachusetts, USA. The population examined in our study was collected from maritime pine in Carnota, Coruña Province, Spain. Females of the Spanish population were characterized by a truncate lip region with two annuli and a slightly protruding labial disc, body annuli more than 300, lateral fields marked by breaks or anastomoses of body annuli, without longitudinal lines, stylet knobs thick, rounded with a moderate cavity (1–1.5 μm), vulval lips not modified, and an elongate conoid tail with offset terminal region. Morphology and morphometrics of this population were similar to those of the original description ( Reed & Jenkins, 1963), except for minor intraspecific differences including a slightly shorter stylet (84–96 vs. 95–112 μm), smaller b ratio (6.3–7.4 vs. 5.9–8.7) and larger c ratio (9.4–11.1 vs. 14.9– 16.7). The present record of H. vaccinii is the first for Europe and Spain, and the fourth after the report from Iran ( Chenari Bouket, Niknam & Eskandari, 2010). This species was also found on highbush blueberries from Ottawa County, Michigan, USA ( Knobloch & Bird, 1981).

Distinguishing characters of this species include a truncate lip region with two or three annuli, lateral fields marked mostly by breaks of striae, an elongate conoid tail tapering to a distal offset and narrower elongated portion with a narrowly rounded terminus, and the absence of males. Hemicycliophora vaccinii is differentiated from the morphologically similar species Hemicycliophora nucleata Loof, 1968 , H. zuckermani , Hemicycliophora mettleri Jenkin & Reed, 1964 , and H. similis ( Chitambar & Subbotin, 2014) .

HEMICYCLIOPHORA WYEI CORDERO LÓPEZ,

Figures

Figure 1. Photomicrographs of specimens of populations of selected Hemicycliophora species. A–F, anterior region; G–L, lateral field; M–R, posterior region. A, G, M, Hemicycliophora conida (Washington State, USA); B, H, N, Hemicycliophora sp. 3 (Arizona, USA); C, I, O, Hemicycliophora sp. 8 (California, USA); D, J, P, Hemicycliophora raskii (California, USA); E, K, Q, Hemicycliophora sp. 10 (California, USA); F, L, R, Hemicycliophora californica (California, USA). Scale bars: A–F, M–R = 10 μm; G–L = 5 μm.

Figure 2. Photomicrographs of specimens of populations of selected Hemicycliophora species. A–E, anterior region; F–J, lateral field: K–O, posterior region. A, F, K, Hemicycliophora floridensis (topotype, Florida, USA); B, G, L, Hemicycliophora poranga (California, USA); C, H, M, Hemicycliophora sp. 11 (Florida, USA), D, I, N, Hemicycliophora sp. 4 (North Carolina, USA); E, J, O, Hemicycliophora wyei (North Carolina, USA). Scale bars: A–E, K–O = 10 μm; F–J = 5 μm.

Figure 3. Scanning electron microscope (SEM) micrographs of specimens of populations of selected Hemicycliophora species. A–D, lip region; E–L, lateral field. A, Hemicycliophora wyei (North Carolina, USA) (CD683); B, Hemicycliophora poranga (California, USA) (CD714); C, Hemicycliophora sp. 3 (Arizona, USA) (CD715); D, Hemicycliophora californica (California, USA) (CD826B); E, Hemicycliophora gracilis (California, USA) (CD45); F, H. wyei (North Carolina, USA) (CD679); G, H. californica (CD826B); H, H. wyei (CD683); I, H. poranga (CD714); J, Hemicycliophora sp. 3 (CD715); K, Hemicycliophora sp. 4 (North Carolina, USA) (CD675); L, H. californica CD826B). Scale bars: A–C, E, G–J, L = 5 μm; D = 2 μm; F, K = 10 μm.

Figure 4. Photomicrographs of specimens of a new Spanish population of Hemicycliophora iberica Castillo et al., 1989. A, entire female body; B, female pharyngeal region; C, female anterior region; D, posterior region; E, detail of lateral field; F–I, female tail tips. Scale bars: A = 100 μm; B–D, F–I = 20 μm; E = 10 μm. ep, excretory pore.

Figure 5. Photomicrographs of specimens of a Spanish population of Hemicycliophora obtusa Thorne, 1955. A, entire female body; B, female pharyngeal region; C, female anterior region; D, detail of lateral field; E, F, vulval and tail regions; G, pharyngeal region of pre-adult male showing absence of stylet; H, I, detail of spicules and bursa of pre-adult male. Scale bars: A = 100 μm; B, C, E–I = 20 μm; D = 10 μm.

Figure 7. Phylogenetic relationships within populations and species of the genus Hemicycliophora as inferred from Bayesian analysis using the D2-D3 of the 28S rRNA gene sequence data set with the general time reversible substitution model with estimation of invariant sites and assuming a gamma distribution with four categories. Posterior probabilities of over 70% are given for appropriate clades. Newly obtained sequences are indicated in bold.

Figure 9. Ancestral state reconstructions for the genus Hemicycliophora based on parsimony (left maximum parsimony tree) and Bayesian inference (BI; right: BI tree) of A, average body length; B, average stylet length; C, average R (total number of body annuli); D, average RV (number of annuli between posterior end of body and vulva). Posterior probabilities for each character state are indicated as pie charts in the majority consensus BI tree.

Figure 10. Ancestral state reconstructions for the genus Hemicycliophora based on parsimony (left maximum parsimony tree) and Bayesian inference (BI; right: BI tree) of A, vulval lip structure; B, tail shape; C, presence of males. Posterior probabilities for each character state are indicated as pie charts in the majority consensus BI tree.