Alexei V. Tchesunov 7.12 Order Chromadorida Chitwood, 1933

Diagnosis : Chromadorea. Cuticle ornamented with body (cuticle heterogeneous) or the ornamentation may be transverse rows of punctuations as dots, rods or “ basket made up of rods jointed in a “basket weave” . Pharyngos- weave ” and often with lateral differentiation. Anterior toma with dorsal tooth usually larger than ventrosublate- sensilla arranged in two or three circles, i.e., an ante- ral ones; teeth hollow or solid; denticles may be present; rior circle of inner labial sensilla (usually papillae, but three nearly equal solid teeth also occur in some genera. may be setiform), an outer circle of outer labial sensilla Male monorchic with anterior testis (synapomorphy); pre- (seti- or papilliform) and a third circle of cephalic setae cloacal supplements cup-shaped (never tubular), may be (pattern six + six + four); the second and third circles absent. Females with two antidromously reflexed ovaries, may form a single circle of ten sensilla (pattern six + ten). the anterior gonad to the right of the intestine, the pos- In the case of pattern six + six + four, the four cephalic terior gonad to the left of the intestine (synapomorphy). setae are longer than the six outer labial sensilla, and in Mostly marine. Five subfamilies. the case of pattern six + ten, the six outer labial sensilla are longer than the four cephalic setae of the same joint circle. Amphideal fovea variable, simple spiral, comma- 7.12.1.1 Subfamily Chromadorinae like, transverse loop or slit, or multispiral; when spiral, Filipjev, 1917 amphideal fovea usually located posterior to the cepha- lic setae, but sometimes lying between the four cephalic Diagnosis (after Decraemer & Smol 2006): Chromadori- setae and are then difficult to observe. Cephalic helmet dae. Cuticle homo- or heterogeneous, usually without not developed as such, but head cuticle may be slightly or with slight lateral differentiation made by larger dots inflated within several genera. Cheilostoma with twelve (only in Chromadorella, the lateral larger dots arranged in cheilorhabdia (folds or rugae); pharyngostoma of varia- longitudinal rows). Outer labial papilliform sensilla and ble size and shape, very often armed with a single bigger four cephalic setae in two separate circles. Amphideal dorsal tooth and two or more smaller ventrosublateral fovea transverse slit-like, difficult to observe under light teeth or denticles. Pharynx largely evenly muscular, microscopy. Buccal cavity armed with three nearly equal cylindrical, posteriorly often widened or terminated with solid teeth (except for Prochromadora with the dorsal muscular bulb with well-cuticularized luminal walls. tooth single or prevails). Pharyngeal tissue not enlarged Female reproductive system didelphic-amphidelphic with around the buccal cavity. Simple and distinctly defined antidromously reflexed ovaries. Males monorchic or dior- posterior pharyngeal bulb. Preanal cup-shaped supple- chic. Precloacal supplements cup-shaped, fine tubular or mentary organs usually present. setose. Caudal glands and spinneret present. Marine free- Type genus: Chromadora Bastian, 1865 living, a few species epibiotic, uncommon in freshwater and soil. Six families. 7.12.1.1.1 Genus Atrochromadora Wieser, 1959 (Fig. 7.92 A) 7.12.1 Family Chromadoridae Diagnosis : Chromadorinae. Homogeneous р unctated Filipjev, 1917 body cuticle pattern along the body and with lateral diffe- rentiation of larger dots. Amphideal fovea transverse oval, Diagnosis: Chromadorida. Amphideal fovea as simple open loop-shaped. Pharyngostoma with three solid teeth, transverse slit-like, often inconspicuous, or ventrally the dorsal tooth larger than the ventrosublateral teeth. wound spiral, located between the cephalic setae or pos- Males usually with cup-shaped precloacal supplements. terior to them. Cuticular ornamentation as punctuations, Marine. which may be evenly distributed and of equal size (cuticle Number of species: 12 homogeneous), or unevenly distributed, for example, Type species: Atrochromadora parva (de Man, 1893) enlarged in the lateral body regions or different along the Wieser, 1954 ( = Spiliphera parva de Man, 1893)

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B A

H

D

C

E

F

G

I

Fig. 7.92 : Examples of Chromadorinae. A, Atrochromadora obscura (from Wieser 1959, Fig. 66, A – C); B, Chromadora yamadai (from Kito 1978, Fig. 2); C, Chromadorella edmondsoni (from Wieser 1959, Fig. 67 A – D); D, Chromadorina astacicola (from Wieser 1968, Abb. 1); E, Prochromadora oerleji (head and anterior body) and P. erythrophthalma (male tail) (from Gerlach 1951, Abb. 21, 22); F, Prochromadorella obtusidens (from de Coninck & Stekhoven 1933, Fig. 56 – 57); G, H, Punctodora exochopora (from Hopper 1963, Fig. 1 – 3, 5, 9); I, Timmia acuticauda (from Gal’ tsova 1985, Fig. 19).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.1 Family Chromadoridae Filipjev, 1917 375

7.12.1.1.2 Genus Chromadora Bastian, 1865 by a small elevation of the buccal wall or small ventro- (Fig. 7.92 B) sublateral teeth. Precloacal supplements present. Seven marine and one freshwater species. Diagnosis : Chromadorinae. Homogeneous punctated body Number of species: 8 cuticle pattern along the body, with lateral differentiation Type species: Prochromadora megodonta Filipjev, 1922 of larger dots. Transverse slit-like amphideal fovea. Pha- ryngostoma with three solid teeth, the dorsal tooth larger than the ventrosublateral teeth. Ocelli may be present. 7.12.1.1.6 Genus Prochromadorella Micoletzky, 1924 Males usually with cup-shaped precloacal supplements. (Fig. 7.92 F) Differs from very close Atrochromadora by only amphid shape, but the differentiation is not always clear. Marine. ( = Trichromadora Kreis, 1929) Number of species: 31 Diagnosis (after Decraemer & Smol 2006): Chromado- Type species: Chromadora nudicapitata Bastian, 1865 rinae. Heterogeneous punctated body cuticle along the body, lateral dots may be enlarged but not arranged in longitudinal rows. Amphideal fovea transversely oval, 7.12.1.1.3 Genus Chromadorella Filipjev, 1918 obscure, located between the four cephalic setae. Buccal (Fig. 7.92 C) cavity with three solid teeth of about equal size or the two ventrosublateral teeth smaller. Ocelli may be present. Diagnosis: Chromadorinae. Heterogeneous punctated Males with cup-shaped precloacal supplements. Marine. body cuticle along the body, with lateral differentiation in Number of species: 35 longitudinal rows of larger dots. Amphideal fovea trans- Type species: Prochromadorella neapolitana (de Man, verse oval in shape and slightly bent. Pharyngostoma with 1876) Micoletzky, 1924 ( = Chromadora neapolitana de three solid teeth of about equal size. Posterior pharyngeal Man, 1876) bulb not distinctly set off and with plasmatic interruptions that may appear double in some species. Males with five to twelve (mostly five) cup-shaped precloacal supplements. 7.12.1.1.7 Genus Punctodora Filipjev, 1929, Differs from the similar Neochromadora by only structure (Fig. 7.92 G, H) of the pharyngostomal teeth. Marine. Number of species: 16 Diagnosis : Chromadorinae. Cuticle finely striated and finely Type species: Chromadorella filiformis (Bastian, 1865) homogeneously punctated along the body; lateral dots, Filipjev, 1918 ( = Chromadora filiformis Bastian, 1865) some bigger than submedian dots. Amphideal fovea flatte- ned, spiral and situated at the level of dorsal tooth. Stoma with well-developed dorsal tooth and two smaller ventro- 7.12.1.1.4 Genus Chromadorina Filipjev, 1918 sublateral teeth visible at the anterior and posterior border (Fig. 7.92 D) of an indentation, also described as two transverse folds. Ocelli present. Secretory-excretory in head region. Well- Diagnosis: Chromadorinae. Homogeneous cuticle with defined posterior pharyngeal bulb. From one to 18 cup- transverse rows of dots but without obvious lateral differen- shaped preanal supplements. Three species are freshwater, tiation. Amphideal fovea when visible, transverse slit-like. one is brackish and another is freshwater and brackish. Pharyngostoma with three nearly equal solid teeth. Ocelli Number of species: 5 may be present. Cup-shaped precloacal supplements present. Type species: Punctodora ratzeburgensis (Linstow, 1876) Marine and brackish species and four freshwater species. Filipjev, 1929 ( = Chromadora ratzeburgensis Linstow, 1876 Number of species: 26 Type species: Chromadorina obtusa Filipjev, 1918 7.12.1.1.8 Genus Timmia Hopper, 1961 (Fig. 7.92 I)

7.12.1.1.5 Genus Prochromadora Filipjev, 1922 ( = Parachromadora Timm, 1952, homonym to Parachro- (Fig. 7.92 E) madora Micoletzky, 1913) Diagnosis: Chromadorinae. Homogeneous cuticle with Diagnosis: Chromadorinae. Cuticle with homogeneous transverse rows of dots but without lateral differentiation. punctuation, without lateral differentiation. A single large Amphideal fovea slit-like, when visible. Buccal cavity dorsal tooth opposed by a ventrosublateral pit or at most with three teeth of about equal size. A tubular curved

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 376 7.12 Order Chromadorida Chitwood, 1933 supplement immediately preanal in addition to the 7.12.1.2.3 Genus Austranema Inglis, 1969 (Fig. 7.93 C) typical cup-shaped pre-cloacal supplements. Timmia is similar to Chromadorina except for the tubular pre-cloacal Diagnosis : (after Inglis 1969): Euchromadorinae. Cuticle supplement. Marine. complex, marked dorsally and ventrally, and thickened over Number of species: 3 pharyngeal region. Six outer labial and four cephalic setae Type species: T. parva (Timm, 1952) Hopper, 1961 arranged in a common circle. Dorsal tooth generally hollow. ( = Parachromadora parva Timm, 1952) Pharynx evenly expanding posteriorly. In males, precloacal midventral modification of cuticle present. Tail long and thin. Number of species: 5 7.12.1.2 Subfamily Euchromadorinae Type species: Austranema colesi (Inglis, 1968) Inglis, 1969 Gerlach & Riemann, 1973 ( = Euchromadora colesi Inglis, 1968)

Diagnosis : (after Decraemer & Smol 2006): Chromadoridae. Body cuticle with complex heterogeneous ornamentation 7.12.1.2.4 Genus Crestanema Pastor de Ward, 1985 along the body (rarely homogeneous as in Endeolophos (Fig. 7.93 D) Boucher, 1976), often with lateral differentiation. The six outer labial and four cephalic sensilla setiform may be Diagnosis: (after Pastor de Ward 1985): Euchromadorinae. arranged in a single circle. Amphideal fovea transverse Six outer labial and four cephalic setae arranged in one slit-like or oval, located posterior to the cephalic setae. crown. Buccal armature formed by one dorsal and two Buccal cavity with large dorsal tooth, with or without den- small subventral teeth. Amphideal fovea transversally ticles or smaller ventrosublateral teeth. Pharynx with or oval, with slightly concave posterior margin. Cuticle without defined terminal bulb. Gubernaculum usually with formed by rings with internal anterior and posterior hammer- or L-shaped lateral pieces (wrongly indicated as processes, with alternate hooking. Wing type lateral telamon). Precloacal supplements absent, but a precloacal differentiation nerve ring onward. Spicules simple in differentiation of body cuticle may be present. Marine. structure, gubernaculums with two central pieces and two Type genus: Euchromadora de Man, 1886 lateral guiding pieces (telamon). Type and only species: Crestanema patagonicum Pastor de Ward, 1985 7.12.1.2.1 Genus Actinonema Cobb, 1920 (Fig. 7.93 A)

( = Pareuchromadora Stekhoven & Adam, 1931) 7.12.1.2.5 Genus Endeolophos Boucher, 1976 Diagnosis : Euchromadorinae. Cuticle heterogeneous with (Fig. 7.93 E) lateral differentiation as a ridge beginning at the end of the pharynx. Six outer labial and four cephalic setae Diagnosis : Euchromadorinae. Cuticle heterogeneous, orna- arranged in one circle of ten setae. Amphideal fovea con- mented with longitudinal stick-like elements, with lateral spicuous transversally oval with a double contour. Poste- differentiation but not as a ridge. Six outer labial and four rior pharyngeal bulb may be developed or not. Spicules cephalic sensilla arranged in two separate crowns of setae. simple, curved, with L-shaped telamon. Amphideal fovea as a transversal slit at the level of the Number of species: 7 cephalic setae. Buccal cavity with dorsal tooth and two tiny Type species: A. pachydermatum Cobb, 1920 subventral teeth. No posterior pharyngeal bulb developed. Copulatory apparatus without lateral pieces (telamons). Number of species: 5 7.12.1.2.2 Genus Adeuchromadora Boucher & Type species: Endeolophos fossiferus (Wieser, 1954) de Bov é e, 1971 (Fig. 7.93 B) Boucher, 1976 ( = Nygmatonchus fossiferus Wieser, 1954)

Diagnosis: (after Boucher & de Bové e 1971): Euchroma- dorinae. Cuticle heterogeneous, with lateral ridge along 7.12.1.2.6 Genus Euchromadora De Man, 1886 the body. Amphideal fovea large, transversally elliptical. (Fig. 7.93 F) Buccal cavity armed with a pointed dorsal tooth and two small subventral teeth. Diagnosis: Euchromadorinae. Complex cuticle heterogene- Type and only species: Adeuchromadora megamphida ous, structured with hexagonal or ovoid punctations ante- Boucher & de Bov é e, 1971 riorly and posteriorly, with slimmer markings restricted

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A B C

F E D

I

J G

L K

H

Fig. 7.93 : Examples of Euchromadorinae. A, Actinonema celtica (from Boucher 1976, Fig. 1); B, Adeuchromadora megamphida (from Boucher & de Bov ée 1971, Fig. 2); C, Austranema alii (from Murphy 1965, Fig.1, 2); D, Crestanema patagonicum (from Pastor de Ward 1985, Fig. 7 H – J); E, Endeolophos spinosos galapagensis (head) and E. minutus (male tail) (from Blome 1982, Abb. 28, 29); F, Euchromadora vulgaris head (from Coles, 1965, Fig. 10) and E. ezoensis male tail (from Kito 1977, Fig. 1, 10); G, Graphonema northumbriae (from Warwick & Coles 1975, Fig. 60); H, Parapinnanema harveyi (from Warwick & Coles 1975, Fig. 62); I, Protochromadora parafricana (from Gerlach 1958, Abb. 4); J, Rhips paraornata (from Platt & Zhang 1982, Fig. 9); K, Steineridora borealis (from Kito 1977, Fig. 2); L, Trochamus carinatus (from Boucher & de Bov é e 1971, Fig. 1).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 378 7.12 Order Chromadorida Chitwood, 1933 to the lateral surface over the middle of the body. Trans- fovea as a transversal slit without a cuticular fringe. versally elliptical amphideal fovea without surrounding Dorsal tooth larger than two subventral ones. No posterior cuticular fringe. Four cephalic setae. Buccal cavity with pharyngeal bulb developed. No precloacal modification of large dorsal tooth, subventral teeth and rows of denticles. the midventral cuticle. Tail short and stout. No distinct posterior pharyngeal bulb. Gubernaculum with Number of species: 3 prominent hammer- or L-shaped lateral pieces (telamons). Type species: Protochromadora scampae (Coles, 1965) Number of species: 16 Inglis 1969 ( = Euchromadora scampae Coles, 1965) Type species: Euchromadora vulgaris (Bastian, 1865) de Man, 1886 ( = Chromadora vulgaris Bastian, 1865) 7.12.1.2.10 Genus Rhips Cobb, 1920 (Fig. 7.93 J)

7.12.1.2.7 Genus Graphonema Cobb, 1898 (Fig. 7.93 G) Diagnosis : Euchromadorinae. Cuticle heterogeneous with lateral differentiation as a narrow ridge beginning at the Diagnosis : Euchromadorinae. Complex cuticle heterogene- end of the pharynx. Six outer labial and four cephalic ous, structured with hexagonal or ovoid punctations ante- setae arranged in a common circle. Amphideal fovea cons- riorly and posteriorly, with slimmer markings restricted to picuous transversally oval with a double contour. Pharynx the lateral surface over the middle of the body. Four cepha- gradually enlarged posteriorly. Spicules double-jointed. lic setae. Transversally elliptical amphideal fovea with Gubernaculum with lateral pieces. cuticular fringe. Buccal cavity with large, seemingly hollow Number of species: 7 dorsal tooth, subventral teeth; rows of denticles absent. Type species: Rhips ornata Cobb, 1920 No distinct posterior pharyngeal bulb. Gubernaculum with prominent hammer- or L-shaped lateral pieces (telamons). Number of species: 6 7.12.1.2.11 Genus Steineridora Inglis, 1969 (Fig. 7.93 K) Type species: Graphonema vulgare Cobb, 1898 Diagnosis (after Inglis 1969): Euchromadorinae. Massive squarish dorsal onchium and sickle-like prominent 7.12.1.2.8 Genus Parapinnanema Inglis, 1969 onchia laterally and ventrally. Posterior pharyngeal bulb (Fig. 7.93 H) present. No precloacal cuticular modification. Number of species: 4 Diagnosis : Euchromadorinae. Cuticle usually sculptured Type species: Steineridora loricata (Steiner, 1916) Inglis, in minute hexagonal blocks that become more elongate 1969 ( = Spiliphera loricata Steiner, 1916) on posterior part of the body; cuticle with punctations dorsally and ventrally; cuticle very thick over pharyn- geal region and battlement-like. Six outer labial and four 7.12.1.2.12 Genus Trochamus Boucher & de Bov é e, cephalic setae arranged in a common circle. Transversally 1971 (Fig. 7.93 L) elliptical amphideal fovea with surrounding cuticular fringe. Buccal armament consists of larger dorsal tooth Diagnosis (after Boucher & de Bové e 1971: Euchromadori- and two smaller subventral teeth; no additional denticles. nae. Body cuticle heterogeneous, with a crest-like lateral No posterior pharyngeal bulb. In males, ventral cuticle differentiation. Six outer labial and four cephalic setae anterior to the cloaca forms a prominent modification. arranged in one crown. Amphideal fovea as poorly visible Tail long and slim. transversal slit. Buccal cavity armed with one dorsal tooth Number of species: 6 and two small subventral denticles. No lateral pieces at Type species: Parapinnanema wilsoni Inglis, 1969 spicular apparatus. Number of species: 3 Type species: Trochamus carinatus Boucher & Bové e, 1972 7.12.1.2.9 Genus Protochromadora Inglis, 1969 (Fig. 7.93 I) 7.12.1.3 Subfamily Hypodontolaiminae Diagnosis (after Inglis 1969): Euchromadorinae. Cuticle de Coninck, 1965 complex, without punctations dorsally and ventrally, not markedly thick over pharyngeal region. Cephalic setae Diagnosis: Chromadoridae. Cuticle punctated hetero- sensilla separated as a crown of four setae. Amphideal geneously or rarely homogeneously, with or without

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.1 Family Chromadoridae Filipjev, 1917 379 lateral differentiation. Six outer labial papillae (except 7.12.1.3.3 Genus Deltanema Kreis, 1929 Megodontolaimus with outer labial setae) and four cepha- lic setae in two separate circles. Amphideal fovea distinct Diagnosis (after Decraemer & Smol 2006): Hypodonto- or rather obscure, transverse flattened oval, generally laiminae. Body cuticle with fine transverse annulations; located between the four cephalic setae. Stoma funnel- ornamentation not described. Amphideal fovea trans- shaped, armed with hollow teeth; the large dorsal tooth verse slit-like, located anterior to a large dorsal tooth. may be opposed by two smaller ventrosublateral teeth, Buccal cavity large, with well-developed dorsal tooth and denticles may be present; anterior part of pharynx often two (?) smaller ventrosublateral teeth. with prominent dorsal muscular swelling. Males with Type and only species: Deltanema parvum Kreis, 1929 cup-like precloacal supplements, rarely absent. Marine, Marine; genus originally described as close to Chro- brackish water. madorina and Chromadorita but distinguished from Type genus: Hypodontolaimus de Man, 1886 these genera by well-developed teeth. Deltanema is a monotypic genus known only from females of the type population. The genus was synonymized with Metalin- 7.12.1.3.1 Genus Chromadorissa Filipjev, 1917 homoeus by Hope and Murphy (1972) but recognized as (Fig. 7.94 A) valid by Lorenzen (1981, 1994). The limited description and illustrations of the type species are responsible for Diagnosis (after Filipjev 1917): Hypodontolaiminae. the unclear taxonomic status of this marine genus. Here, Cuticle with heterogeneous punctated ornamentation and the genus Deltanema is considered as genus dubius. lateral differentiation of larger dots arranged in two longi- tudinal rows. Buccal cavity with one large hollow dorsal 7.12.1.3.4 Genus Denticulella Cobb, 1933 (Fig. 7.94 C) tooth and smaller ventrosublateral teeth. Peribuccal pha- ryngeal tissue swollen moderately and nearly symmetri- Diagnosis (after Dashchenko 2002): Hypodontolaiminae. cally. Pharyngeal posterior bulb bipartite. Males with five Body cuticle with heterogeneous punctation along the to six complex supplements. Marine and brackish-water body, lateral differentiation of larger dots not arranged in species. longitudinal rows and, in the region of the buccal cavity, Number of species: 3 a pattern recalling the plaiting of a basketball may be Type species: Chromadorissa beklemishevi Filipjev, 1917 present. Buccal cavity with a hollow dorsal tooth, two smaller ventrosublateral teeth and numerous additional denticles. Pharynx anteriorly with asymmetrical dorsal 7.12.1.3.2 Genus Chromadorita Filipjev, 1922 swelling with a degree of swelling related to the size of (Fig. 7.94 B) dorsal tooth, and posteriorly with a single end bulb. Males with cup-shaped precloacal supplements. Marine. ( = Allgeniella Strand, 1974) Number of species: 2 Diagnosis : Hypodontolaiminae. Cuticle ornamentation Type species: Denticulella pellucida Cobb, 1933 homogeneous exceptionally with slightly more pro- The genus is intermediate in position between the nounced punctation at the level of lateral fields. Hollow genera Chromadorita and Dichromadora , (Decraemer & teeth, mostly one dorsal and one or two ventrosublateral, Smol, 2006) the cuticle being laterally differentiated into rarely one indistinct dorsal tooth only; sometimes tiny larger dots, not arranged into longitudinal rows as in Chro- denticles may be present. Pharynx may be swollen ante- madorita , and the buccal teeth similar in arrangement as riorly in various degrees; posterior bulb single. Precloa- in Dichromadora; denticles considered as typical of the cal supplements present or absent. Mostly marine, two genus were also observed in Dichromadora hyalocheile species were also recorded in fresh water. Gerlach, 1951. Wieser (1954) transferred Dichromadora Number of species: 27 stygia Gerlach, 1952 to Denticulella , but Blome (1974) trans- Type species: Chromadorita demaniana Filipjev, 1922 ferred it to a new genus Parachromadorita Blome, 1974. A sub-group within the Innocuonema - Chromadorita complex lacks precloacal supplements, has a somewhat elongated angular punctation and a tail with the tip bent 7.12.1.3.5 Genus Dichromadora Kreis, 1929 to the left and dorsally curved. Platt and Warwick (1988) (Fig. 7.94 D) remarked that some confusion may arise between Chro- madorita and Innocuonema Inglis, 1969; both genera need Diagnosis: Hypodontolaiminae. Cuticle ornamented revision. homogeneously with a pronounced lateral differentiation

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D

C A B

E F G

J

H I

K L M

Fig. 7.94 : Examples of Hypodontolaiminae. A, Chromadorissa beklemishevi (adapted from Filipjev 1917, Fig. A – C, G); B, Chromadorita tenuis (from Gerlach 1965, Abb. 23); C, Denticulella boreala (from Dashchenko 2002, Fig. 2); D, Dichromadora geophila (from Gerlach 1951, Abb. 9); E, Hypodontolaimus colesi (from Platt & Warwick 1988, Fig. 73); F, Innocuonema flaccidum (from Wieser 1959, Fig. 56); G, Karkinochromadora lorenzeni (from Blome 1982, Abb. 26 I – K); H, Megodontolaimus coxbazari (from Timm 1969, Fig. 1 – 2); I, Neochromadora poecilosoma (from Gerlach 1951, Abb. 9); J, Panduripharynx ornata (from Timm 1961, Fig. 29, anterior body) and P. pacifica (from Belogurov et al. 1985, Fig. 2, head); K, Parachromadorita stygia (from Blome 1974, Abb. 6 – 14); L, Ptycholaimellus adocius (from Dashchenko & Belogurov 1984, Fig. 2, head, optical section), P.E hibernus (from Eskin & Hopper 1985, Fig. 4 – 5, surface view of head, male tail) and P. pandispiculatus (from Hopper 1961, Fig. 2, pharyngeal region); M, Spilophorella candida (from Gerlach 1951, Abb. 1, head and pharyngeal region) and S. paradoxa (from Gerlach 1951, Abb. 2).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.1 Family Chromadoridae Filipjev, 1917 381 in the form of two longitudinal rows of enlarged dots longitudinal rows of enlarged dots. Amphideal fovea bordering the lateral field without ornamentation. Buccal rounded loop-shaped. Buccal cavity with two conse- cavity armed either with a single S-shaped hollow dorsal cutive dorsal teeth: a small anterior and a larger pos- tooth or with three teeth, the dorsal one being larger than terior and two small ventrosublateral teeth. Peribuccal the two ventrosublateral ones; sometimes denticles may pharyngeal tissue swollen dorsally. End bulb distinct, be present. Pharynx usually not swollen anteriorly; pos- pear-shaped. Weak precloacal supplements present terior bulb single. Precloacal supplements in male absent with cyatholaimid arrangement, i.e., with grouped or present. Marine, with three brackish water species. distribution. Number of species: 25 Type and only species: Karkinochromadora lorenzeni Type species: Dichromadora cephalata (Steiner, 1916) (Jensen, 1980) Blome, 1982 ( = Chromadora lorenzeni Kreis, 1929 ( = Chromadora cephalata Steiner, 1916) Jensen, 1980) Originally, the Karkinochromadora type species was described as a Chromadora species (Jensen, 1980) with 7.12.1.3.6 Genus Hypodontolaimus de Man, 1886 a single dorsal tooth; the anteriormost dorsal tooth was (Fig. 7.94 E) apparently interpreted as part of the folded stoma wall.

( = Iotadorus Cobb, 1920) Diagnosis: Hypodontolaiminae. Body cuticle with puncta- 7.12.1.3.9 Genus Megodontolaimus Timm, 1969 ted ornamentation of two longitudinal rows of larger dots. (Fig. 7.94 H) Buccal cavity with large S-shaped hollow dorsal tooth and a dorsal apophysis. Pharynx swollen in buccal region to Diagnosis: Hypodontolaiminae. Body cuticle with larger accommodate the dorsal tooth; end bulb single. Precloa- dots toward the extremities of the body or only anteri- cal supplements usually present, cup-shaped. Marine and orly, and with lateral differentiation of larger dots at the brackish. borders of the lateral field; two or six alae are present, Number of species: 27 showing fine cross bars. Inner and outer labial sensilla as Type species: Hypodontolaimus inaequalis (Bastian, 1865) short setae, cephalic sensilla as much longer setae. Buccal de Man, 1886 ( = Spiliphera inaequalis Bastian, 1865) cavity with a large hollow ventral tooth with hooked tip Hypodontolaimus is similar to Dichromadora except for and a two-pronged dorsal tooth with crescent thickening shape of the dorsal tooth and apophysis. along the buccal wall. Posterior pharyngeal bulb double. No precloacal supplements. Gubernaculum with lateral 7.12.1.3.7 Genus Innocuonema Inglis, 1969 sleeve. ( Fig. 7.94 F) Number of species: 2 Type species: Megodontolaimus coxbazari Timm, 1969 Diagnosis: Hypodontolaiminae. Body cuticle with complex and obvious punctations, no lateral differenti- ation. Buccal cavity with large hollow dorsal tooth and 7.12.1.3.10 Genus Neochromadora Micoletzky, 1924 associated slight dorsal swelling of the pharynx in this (Fig. 7.94 I) region. Posterior bulb definite, single. Precloacal supple- ments absent. Marine. Diagnosis : Hypodontolaiminae. Body cuticle ornamenta- Number of valid species: 6 tion complex, with lateral differentiation visible as two or Type species: Innocuonema flaccidum (Wieser, 1959) three longitudinal rows of large dots. Buccal cavity with Inglis, 1969 ( = Graphonema flaccidum Wieser, 1959) a hollow dorsal tooth and two ventrosublateral teeth. A revision of the genus is recommended because the Pharynx anteriorly enlarged, especially in association with difference between the genera Chromadorita and Innocuo- the dorsal tooth and posteriorly with a well-developed end- nema is not quite clear. bulb. Posterior pharyngeal bulb well-developed, single. Male usually with numerous cup-shaped precloacal sup- plements. Most species marine, one species brackish and 7.12.1.3.8 Genus Karkinochromadora Blome, 1982 another one freshwater. (Fig. 7.94 G) Number of species: 31 Type species: Neochromadora poecilosoma (de Man, 1893) Diagnosis (modified after Blome 1982): Hypodontolai- Micoletzky, 1924 ( = Chromadora poecilosoma de Man, minae. Body cuticle with lateral differentiation of two 1893)

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7.12.1.3.11 Genus Panduripharynx Timm, 1969 7.12.1.3.14 Genus Spilophorella Filipjev, 1917 (Fig. 7.94 J) (Fig. 7.94 M)

Diagnosis: Hypodontolaiminae. Cuticle ornamentation Diagnosis: Hypodontolaiminae. Cuticle with complex complex; lateral differentiation with larger dots bordering heterogeneous punctated ornamentation with lateral the lateral field (alae). Stoma broad, with well-cuticularized differentiation of larger dots arranged in longitudinal walls, large hollow dorsal tooth with small apophysis, two rows. Deep buccal cavity with a long hollow dorsal tooth. smaller hollow ventrosublateral teeth, and solid denticles. Peribuccal pharyngeal tissue swollen slightly and nearly Peribuccal pharyngeal tissue swollen dorsally; posteror symmetrical. Pharynx with elongated double posterior bulb double, massive, panduriform, with heavy internal bulb. Tail terminated with a long pointed caudal tube sclerotizations. No precloacal supplements. ( spinneret). Mostly marine, some species rarely were Number of species: 3 recorded in brackish habitats. Type species: Panduripharynx ornata Timm, 1952 Number of species: 12 Type species: Spilophorella paradoxa (de Man, 1888) Filipjev, 1917 ( = Spiliphera paradoxa de Man, 1888) 7.12.1.3.12 Genus Parachromadorita Blome, 1974 (Fig. 7.94 K) 7.12.1.4 Subfamily Harpagonchinae Diagnosis (after Blome, 1974): Hypodontolaiminae. Body Platonova & Potin, 1972 cuticle with lateral differentiation of irregular larger dots not arranged into longitudinal rows. Buccal cavity with a Diagnosis : Chromadoridae. Body cuticle with homogene- large hollow dorsal tooth, two smaller ventrosublateral ous punctuation; no lateral differentiation. A circle of six teeth and a field of denticles. Peribuccal tissue swollen setae within anterior sensilla pattern. Buccal cavity with dorsally; pharyngeal end bulb single, pear-shaped. Male three movable triangular-shaped mandibles with ante- with cup-shaped precloacal supplements. rior solid hooks. Pharynx enlarged anteriorly around the Type and only species: Parachromadorita stygia (Gerlach, mandibular apparatus and posteriorly widened. Precloa- 1952) Blome, 1974 ( = Dichromadora stygia Gerlach, 1952) cal supplements present (cup-shaped) or absent. Marine, The genus is differentiated from the related genera ectosymbionts of Polychaeta. Chromadorita, Dichromadora and Chromadora by the type Type genus: Harpagonchus Platonova & Potin 1972 of lateral differentiation of the cuticle ornamentation as well as the shape of the amphideal fovea in the first two genera. It is also differentiated by the shape of the amphi- 7.12.1.4.1 Genus Harpagonchus Platonova & Potin, 1972 deal fovea from the genus Denticulella that also possesses (Fig. 7.95 A – D) a field of denticles. Marine. Diagnosis : Harpagonchinae. Pharynx posteriorly widened but not forming a true terminal bulb. Cup-shaped midven- 7.12.1.3.13 Genus Ptycholaimellus Cobb, 1920 tral precloacal supplementary organs present. (Fig. 7.94 L) Number of species: 2 Type species: Harpagonchus averincevi Platonova & Potin, Diagnosis : Hypodontolaiminae. Body cuticle with punc- 1972 tated ornamentation of two longitudinal rows of larger dots. Labial region usually offset and narrower than the rest of the head. Dorsal tooth in the buccal cavity large, 7.12.1.4.2 Genus Harpagonchoides Platonova & hollow, with a dorsal apophysis. Peribuccal pharyngeal Potin, 1972 tissue swollen dorsally to accommodate the dorsal tooth. There, a pair of small globular cuticular bodies may be Diagnosis : Harpagonchinae. Pharynx posteriorly terminated present on the outside of the peribuccal swelling. Pos- with a bulb. No precloacal supplementary organs in males. terior pharyngeal bulb double. Precloacal supplements Type and only species: Harpagonchoides crassus mainly absent. Platonova & Potin, 1972. Number of species: 15 Platonova and Potin (1972) initially described all Type species: Ptycholaimellus carinatus Cobb, 1920 the species and established the family Harpagonchidae

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L F G E

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Fig. 7.95 : Examples of Harpagonchinae and Spilipherinae. A – D, Harpagonchus averincevi (from Platonova & Potin 1972, Fig. 1); E – L, Structures of Acantholaimus spp. E – J: A. ewensis , details (from Platt & Zhang 1982, Fig. 7); K, Spermatozoon of A. maks (from Gerlach et al. 1979, Abb. 2F); L, Spermatozoon of A. calathus (from Gerlach et al. 1979, Abb. 8G); M, Spiliphera gracilicauda (from Platt & Warwick 1988, Fig. 40).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 384 7.12 Order Chromadorida Chitwood, 1933 within Chromadorida. Lorenzen (1981, 1994) restudied Acantholaimus distinguishes itself from other genera the type specimens and lowered the rank of Harpagon- of Chromadoridae by some features, particularly by struc- chidae to the level of subfamily within the family Chro- turally complex spermatozoa. The latter may be compa- madoridae, particularly on the basis of a single testis in rable in size with ova in utery; usually, the giant sperma- the males. tozoa are visibly very few in number within a male gonad The harpagonchines are evidently quite peculiar in (e.g., Gerlach et al. 1979). the mode of life. All the species were collected from sub- Acantholaimus is a typical deep-sea genus occu- antarctic and antarctic polychaete worms, Harpagonchus ring in all oceans in the continental slope and abyssal averincevi and H. similis on the gills of Aglaophamus mac- plains (de Mesel et al. 2006). Acantholaimus is present roura (Nephtyidae), and Harpagonchoides crassus between in nearly every sample of deep-sea sediment, where parapodia of Hemipodus digitifera (Glyceridae). The nema- moreover, the genus may be very abundant and species- todes fasten themselves onto body of polychaetes using rich. Not infrequently, several species occur together in mandibular hooks. The only other known example of such one sample, and then they differ from one another in ectosymbiosis (or even ectoparasitism) of nematodes on body size, tail length and details of the buccal arma- polychaetes is Theristus polychaetophilus (Xyalidae) on ment (Miljutina & Miljutin 2012; Miljutina et al. 2013); Scolelepis squamata presented by Hopper (1966). possibly those species can occupy different ecological niches in deep-sea nematode communities.

7.12.1.5 Subfamily Spilipherinae Filipjev, 1918 7.12.1.5.2 Genus Spiliphera Bastian, 1865 (Fig. 7.95 M)

Diagnosis (after Decraemer & Smol 2006): Chromadoridae. ( = Statenia Allg é n, 1930) Body cuticle with homogeneous or heterogeneous puncta- Diagnosis: Spilipherinae. Amphideal fovea in shape of tion, with or without lateral differentiation. Buccal cavity clear open-looped spiral. Spicules not expanded proxi- with three or more solid teeth with or without apophyses. mally. Tail rather long, filiform. Six outer labial setae and four cephalic setae usually inser- Number of valid species: 2 ted at the same level. Amphideal fovea situated laterally Type species: Spiliphera elegans Bastian, 1865 on the head, spiral, i.e., either cryptospiral with a circu- lar outline or a single-loop spiral with at most 1.5 turns. Pharynx with subdivided end bulb. Precloacal supple- 7.12.2 Family Cyatholaimidae ments setose or absent. Tail conical or elongate. Marine. Filipjev, 1918 Type genus: Spiliphera Bastian, 1865

Diagnosis : Chromadorida. Cuticle with transverse rows of punctations. Lateral punctations may be larger, irregu- 7.12.1.5.1 Genus Acantholaimus Allg é n, 1933 lar or arranged in longitudinal rows. Inner labial sensilla (Fig. 7.95 E – L) often setiform; six outer labial sensilla and four cepha- lic sensilla jointed in a single circle of ten setae (with very ( = Neochropmadorina Kreis, 1963) rare exceptions where in separate circles); the six outer Diagnosis: Spilipherinae. Body cuticle punctated with dots labial setae longer than the four cephalic setae. Amphideal variously, often with transverse rows; lateral differentia- fovea lateral, multispiral. Cheilostoma with distinctly cuti- tion may be present as enlarged dots, irregular or arranged cularized twelve rugae. Pharyngostoma with a large dorsal in rows. Amphideal fovea large, round with interrupted tooth, and usually with one or two pairs of smaller ventrosu- posterior rim or nearly comma-shaped. Buccal armament blateral teeth. Pharynx without or rarely with terminal bulb. as three or more solid teeth, minute or large, often protru- Female didelphic-amphidelphic with anterior and posterior sible. Spicules of a peculiar shape, flattened and strongly antidromously reflexed gonads always on different sides of broadened proximally. Spermatozoa giant, pear-shaped, the intestine. Male with two testes situated on opposite sides often structurally complex. No precloacal supplements. of the intestine, rarely with one testis; different types of pre- Tail rather long, filiform. Marine, mostly deep-sea. cloacal supplements or supplements absent. Mostly marine, Number of species: 47 some species in brackish or freshwaters. Type species: Acantholaimus longisetosus Allg é n, 1933 Type genus: Cyatholaimus Bastian, 1865

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7.12.2.1 Subfamily Cyatholaiminae 7.12.2.1.4 Genus Metacyatholaimus Stekhoven, Filipjev, 1918 1942 (Fig. 7.96 D)

Diagnosis (after Decraemer & Smol 2006): Cyatholaimidae. Diagnosis: Cyatholaiminae. Lateral differentiation of three Body cuticle with homogeneous punctation, without to five longitudinal rows of enlarged dots. Teeth reduced. lateral differentiation (except in Marylynnia). Precloacal Posterior pharyngeal bulb present. No precloacal supple- supplements absent, except in Praeacanthonchus where mentary organs. Tail conico-cylindrical. tubular and in Marylynnia where cup-shaped; guberna- Number of species: 6 culum unpaired proximally, except in Marylynnia where Type species: Metacyatholaimus hirschi Stekhoven, 1942 paired. Type genus: Cyatholaimus Bastian, 1865 7.12.2.1.5 Genus Paralongicyatholaimus Stekhoven, 1950 (Fig. 7.96 E) 7.12.2.1.1 Genus Cyatholaimus Bastian, 1865 (Fig. 7.96 A) Diagnosis: Cyatholaiminae. No lateral differentiation. Teeth in the buccal cavity reduced or absent. Spicules (syn. Necticonema Marion, 1970) simple. Gubernaculum simple. No precloacal supplemen- Diagnosis: Cyatholaiminae. No lateral differentiation of tary organs. Tail filiform. the cuticle. No precloacal supplements. Gubernaculum Number of species: 4 paired, distally swollen, squarish, with large pointed Type species: Paralongicyatholaimus mastigodes Stekho- teeth. Tail conical. ven, 1950 Number of species: 13 Type species: Cyatholaimus ocellatus Bastian, 1865 7.12.2.1.6 Genus Paramarylynnia Huang & Zhang, 2007 (Fig. 7.96 F) 7.12.2.1.2 Genus Longicyatholaimus Micoletzky, 1924 (Fig. 7.96 B) Diagnosis (after Huang & Zhang 2007): Cyatholaiminae. Body cuticle with transverse rows of even dots; lateral differentia- Diagnosis: Cyatholaiminae. Lateral differentiation of tion absent. Outer labial and cephalic setae in a single circle. cuticle of larger and more wide-spaced dots. Buccal cavity Buccal cavity with a dorsal tooth and paired subventral small, with a dorsal tooth and without subventral teeth. teeth. Gubernaculum large, distally dilated and not dentate. No posterior pharyngeal bulb. Cup-shaped precloacal Precloacal supplements absent. Tail conico-cylindrical. supplements. Gubernaculum distally dentate, paired. Tail Type and onle species: Paramarylynnia subventrosetata with elongate slender flagellum. Huang & Zhang, 2007 Number of species: 9 Type species: Longicyatholaimus longicaudatus (Mico- 7.12.2.1.7 Genus Phyllolaimus Murphy, 1963 (Fig. 7.96 G) letzky, 1924)

Diagnosis: Cyatholaiminae. Cuticle without lateral differen- tiation as longitudinal rows of enlarged dots. Buccal cavity 7.12.2.1.3 Genus Marylynnia Hopper, 1977 (Fig. 7.96 C) cyathiform with prominent, elongate dorsal tooth; additi- onal teeth may be present. Lips with distinctive foliaceous Diagnosis: Cyatholaiminae. Lateral differentiation of processes of labial rugae. No basal pharyngeal bulb. cuticle of larger and widely spaced dots; two types Number of species: 2 of cuticular pores present. Buccal cavity with dorsal Type species: Phyllolaimus tridentatus Murphy, 1963 tooth and two pairs of small subventral teeth. Cup- shaped precloacal supplements, very slightly cuticu- larized. Gubernaculum distally dentate, paired. Tail 7.12.2.1.8 Genus Praeacanthonchus Micoletzky, conico-cylindrical. 1924 (Fig. 7.96 H) Number of species: 16 Type species: Marylynnia annae (Wieser & Hopper, 1967) Diagnosis: Cyatholaiminae. Lateral differentiation of ( = Longicyatholaimus annae Wieser & Hopper, 1967) larger dots. Teeth in the buccal cavity variably developed

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Fig. 7.96 : Examples of Cyatholaiminae 1. A, Cyatholaimus ocellatus (from Wieser 1955, Abb. 2 A – E); B, Longicyatholaimus marilynae (from Hopper 1972, Fig. 17 – 19); C, Marylynnia annae (from Hopper 1972, Fig. 1 – 3); D, Metacyatholaimus spatiosus (from Wieser 1954, Fig. 99); E, Paralongicyatholaimus macramphis (from Lorenzen 1972a, Abb. 8); F, Paramarylynnia subventrosetata (from Huang & Zhang, 2007, Fig. 1); G, Phyllolaimus tridentatus (from Murphy, 1963, Fig. 2 A, C); H, Praeacanthonchius punctatus (from De Coninck, Stekhoven, 1933, Fig. 31, 32); I, Xyzzors fitzgeraldae (from Inglis 1963, 25, 27); J, Acanthonchus duplicatus (from Wieser 1959, Fig. 42); K, Biarmifer laminatus (from Wieser, 1954, Fig. 97); L, Paracanthonchus serratus (from Wieser 1959, Fig. 40).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.2 Family Cyatholaimidae Filipjev, 1918 387 and may be reduced. Tubular precloacal supplements. 7.12.2.2.3 Genus Paracanthonchus Micoletzky, 1924 Gubernaculum distally dentate, proximally single. (Fig. 7.96 L) Number of species: 10 Type species: Praeacanthonchus punctatus (Bastian, 1865) Diagnosis: Paracanthonchinae. Body cuticle with trans- Micoletzky, 1924 ( = Cyatholaimus punctatus Bastian, verse rows of fine dots, which laterally may be slightly larger 1865) or irregularly arranged. Buccal cavity with larger pointed dorsal tooth and smaller ventrosublateral teeth. Guberna- culum proximally paired, distally expanded and dentate. 7.12.2.1.9 Genus Xyzzors Inglis, 1963 (Fig. 7.96 I) Tubular precloacal supplements. Marine and brackish. Number of species: 67 Diagnosis (after Inglis, 1963): Cyatholaiminae. Lateral dif- Type species: Paracanthonchus caecus (Bastian, 1865) ferentiation of cuticle irregular running full length of the Micoletzky, 1924 ( = Cyatholaimus caecus Bastian, 1865) body. Buccal cavity with a very large dorsal tooth. Spicu- les with doubled proximal knobs. Gubernaculum complex distally with only two processes. 7.12.2.2.4 Genus Paracyatholaimoides Gerlach, 1953 Number of species: 3 (Fig. 7.97 A) Type species: Xyzzors fitzgeraldae Inglis, 1963 Diagnosis : Paracanthonchinae. No evident lateral diffe- rentiation in body cuticle pinctations. Cephalic cuticle thi- 7.12.2.2 Subfamily Paracanthonchinae ckened. Amphideal fovea multispiral and tightly coiled, De Coninck, 1965 and larger in males than in females. Precloacal supple- ments small and may be inconspicuous. Marine. Diagnosis (after Decraemer & Smol 2006): Cyatholaimi- Number of species: 4 dae. Homogeneously punctated body, with or without Type species: Paracyatholaimoides multispiralis Gerlach, lateral differentiation in ornamentation. Pharynx without 1953 end bulb. Pre-cloacal supplements variable, but mainly tubular; rarely absent. Gubernaculum paired proximally. Largely marine, rarely limnetic in one genus. 7.12.2.2.5 Genus Paracyatholaimus Micoletzky, 1922 Type genus: Paracanthonchus Micoletzky, 1924 (Fig. 7.97 B)

Diagnosis (modified after Tchesunov, 2008): Paracanthon- 7.12.2.2.1 Genus Acanthonchus Cobb, 1920 chinae. Lateral differentiation hardly developed. Amphidial (Fig. 7.96 J) fovea multispiral. Pharyngostoma armoured with one dis- tinct dorsal tooth, often supplemented with smaller subven- Diagnosis : Paracanthonchinae. Body cuticle mostly tral teeth and occasionally other denticles in the stegostoma. homogeneously punctated, no lateral differentiation. Pre- Precloacal ventromedian supplements as setae-like organs cloacal supplements tubular, anteriormost much larger half inserted into the body. Gubernaculum hardly dilated at than the other. Gubernaculum simple. Marine. the distal end and distally devoid of denticles or serrations. Number of species: 10 Tail conical or with a more or less slender cylindrical distal Type species: Acanthonchus viviparus Cobb, 1920 portion. Most species are marine but there are also a few brackish water, freshwater species, and even in dune sand. Number of species: 24 7.12.2.2.2 Genus Biarmifer Wieser, 1954 (Fig. 7.96 K) Type species: Paracyatholaimus dubiosus (B ü tschli, 1874) Micoletzky, 1922 ( = Cyatholaimus dubiosus Bü tschli, 1874) Diagnosis : Paracanthonchinae. Punctated cuticle without lateral differentiation. Buccal cavity with a dis- tinct dorsal tooth. Male with inconspicuous cup-shaped 7.12.2.3 Subfamily Pomponematinae precloacal supplements. Gubernaculum distally vari- Gerlach & Riemann, 1973 ously structured. Number of species: 3 Diagnosis (after Decraemer & Smol 2006): Cyatholaimi- Type species: Biarmifer cochleatus Wieser, 1954 dae. Body cuticle punctated, with lateral differentiation

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Fig. 7.97 : Examples of Cyatholaiminae and Achromadoridae. A, Paracyatholaimoides multispiralis (from Gerlach 1953c, Abb. 16); B, Paracyatholaimus intermedius (adapted from Gerlach 1953a, Abb. 5); C, Craspodema reflectans (from Gerlach 1964); D, Minolaimus lineatus (from Vitiello 1970, Fig. 25); E, Nannolaimoides decoratus (from Ott, 1972, Fig. 4, 7); F, Parapomponema hastatum (from Ott 1972, Fig. 13, 16 – 18); G, Pomponema elegans (from Lorenzen, 1972b, Abb. 2); H, Propomponema websteri (from Sharma & Vincx 1982, Fig. 26, 30); I, Xenocyatholaimus delamarei (from Gerlach 1953b, Fig. 1); J, Achromadora sedata (from Gagarin 2001, Fig. 1).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.2 Family Cyatholaimidae Filipjev, 1918 389 in ornamentation. Precloacal supplements knob-like or Number of species: 2 flattened, complicated, consisting of several elements; Type species: Parapomponema hastatum Ott, 1872 gubernaculum paired proximally. Type genus: Pomponema Cobb, 1917 7.12.2.3.5 Genus Pomponema Cobb, 1917 (Fig. 7.97 G)

7.12.2.3.1 Genus Craspodema Gerlach, 1956 Diagnosis: Pomponematinae. Body cuticle heterogene- (Fig. 7.97 C) ous, lateral differentiation as longitudinal rows of enlar- ged dots. Cuticle of head region may be thickened. Buccal Diagnosis : Pomponematinae. Lateral differentiation cavity with a big pointed dorsal tooth, lesser subventral of body cuticle very prominent as longitudinal rows of teeth and additional minute denticles. Gubernaculum enlarged punctations with broad lateral fields between with variously structured distal ends. them. Number of species: 30 Number of species: 2 Type species: Pomponema mirabile Cobb, 1917 Type species: Craspodema octogoniata (Gerlach, 1954) Gerlach, 1956 ( = Kraspodema octogoniatum Gerlach, 1954) 7.12.2.3.6 Genus Propomponema Ott, 1972 (Fig. 7.97 H) 7.12.2.3.2 Genus Minolaimus Vitiello, 1970 (Fig. 7.97 D) Diagnosis . Pomponematinae. Body cuticular ornamenta- tion heterogeneous, with alternating rows of dots and slit- Diagnosis : Pomponematinae. Anterior body end conically like markings, no lateral differentiation except for slight narrowed. Lateral differentiation as three longitudinal lateral irregularity on the tail. Gubernaculum with lateral rows of enlarged dots. Amphideal fovea large, multispiral, flanges bearing blunt teeth. situated at a distance from cephalic apex. Buccal cavity Number of species: 2 very small and unarmed. Tail filiform. Type species: Propomponema foeticolum Ott, 1972 Number of species: 2 Synonymization with Pomponema is possible. Type species: Minolaimus lineatus Vitiello, 1970

7.12.2.4 Subfamily Xenocyatholaiminae 7.12.2.3.3 Genus Nannolaimoides Ott, 1972 Gerlach & Riemann, 1973 (Fig. 7.97 E) Diagnosis (after Decraemer & Smol 2006): Cyatholaimidae. Diagnosis: Pomponematinae. Lateral differentiation of Cuticle with transverse striae, only with punctated orna- cuticle irregular. Buccal cavity with small dorsal tooth mentation anterior to the amphideal fovea and on tail; ten and weakly developed ventrolateral projections. Guberna- longitudinal ridges present. Six outer labial setae and four culum with lateral flanges without teeth or denticles. cephalic setae arranged in a single circle, the four cephalic Number of species: 3 setae being shorter than six outer labial setae. Amphideal Type species: Nannolaimoides armatus (Gerlach, 1964) fovea multispiral, located and about one head width from Ott, 1972 ( = Nannolaimus armatus Gerlach, 1964) anterior end. Buccal cavity large, with large pointed dorsal tooth. Precloacal supplements absent; gubernaculum small. Marine. 7.12.2.3.4 Genus Parapomponema Ott, 1972 Type and only genus: Xenocyatholaimus Gerlach, 1953 (Fig. 7.97 F)

Diagnosis (after Ott 1972): Pomponematinae. Cuticular 7.12.2.4.1 Genus Xenocyatholaimus Gerlach, 1953 ornamentation simple, lateral differentiation consis- (Fig. 7.97 I) ting of two rows of larger dots, beginning anterior to the middle of the pharynx. Gubernaculum with L-shaped Diagnosis : see diagnosis of Xenocyatholaiminae. lateral plates; preanal supplements consisting of an outer Type and only species: Xenocyatholaimus delamarei plate and an inner plate, joined by a constriction. Marine. Gerlach, 1953

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7.12.3 Family Achromadoridae enlarged dots. Pattern of anterior sensilla six + six + four, Gerlach & Riemann, 1973 inner labial sensilla papillose, outer labial sensilla papil- lose or setose, cephalic sensilla setose. Amphideal fovea rounded and spirally coiled in one turn, located at the level Diagnosis : Chromadorida. Cuticle with transverse rows of the stoma. Pharyngostoma tubular, with sclerotized walls of fine punctations, without pronounced lateral diffe- and with three nearly equal teeth anteriorly, or subventral rentiation. Six innner labial sensilla papillose. Six outer teeth, some smaller than the dorsal tooth. Pharynx around labial setae and four shorter cephalic setae form a jointed the stoma slightly thickened and offset; terminal thicke- circle, but seldom may be arranged in two separate circles. ning or bulb also present. Two opposed outstretched testes; Amphideal fovea spiral, situated posterior to the cephalic both gonads on different sides of the intestine. Males with setae. Buccal cavity with a distinct dorsal tooth, subvent- ventral row of precloacal cup-shaped supplements; ventral ral teeth small or absent. Pharynx with moderately deve- precloacal seta present. Tail conical, usually rounded tip. loped terminal bulb. Female anterior and posterior gonad Type genus: Ethmolaimus de Man, 1880. Three genera very situated on the same side of the intestine, either both on close to one another. the left or on the right side. Males rare; precloacal supple- ments inconspicuous or absent. Reproduction mostly by parthenogenesis. A purely limnetic-terrestrial family. 7.12.4.1 Genus Ethmolaimus de Man, 1880 Type genus: Achromadora Cobb, 1913 (Fig. 7.98 A) Lorenzen (1981, 1994) concluded that Achromadori- dae is a holophyletic taxon; the holophyly is established Diagnosis: Ethmolaimidae. Head rounded with labial region with two holapomorphies: 1) position of both anterior not offset. Inner labial sensilla papillose, outer labial sensilla and posterior ovaries on the same side of the midgut and papillose or setose, cephalic sensilla setose or papillose. Most 2) prevailing parthenogenetic mode of reproduction. species are freshwater, some occur in brackish biotopes. According to the tree based on small subunit rDNA sequen- Number of species: 13 ces (Holterman et al., 2008), Achromadoridae and Cyatho- Type species: Ethmolaimus pratensis de Man, 1888 laimudae are sister groups to each other; further, they are grouped with Ethmolaimidae and Chromadoridae. Lorenzen (1981, 1994) included also Kreisonema 7.12.4.2 Genus Paraethmolaimus Jensen, Khera, 1969 and Paradoxolaimus Kreis, 1924, which both 1994 (Fig. 7.98 B) were earlier placed within Leptolaimidae. Both genera are poorly studied yet to understand their relationships Diagnosis: Ethmolaimidae. Head with labial region set off. within Chromadorea. All anterior sensilla as papillae. Brackish and marine. Number of species: 2 7.12.3.1 Genus Achromadora Cobb, 1913 Type species: Paraethmolaimus appendixocaudatus Jensen, 1994 (Fig. 7.97 J)

Diagnosis : see diagnosis of Achromadoridae. 7.12.4.3 Genus Trichethmolaimus Platt, Number of species: 24 Type species: Achromadora minima (Cobb, 1891) (junior 1982 (Fig. 7.98 C) synonym of Achromadora ruricola (de Man, 1880) Diagnosis: Ethmolaimidae. Body cuticle with supporting Micoletzky, 1922) rods elongated in head region. Somatic setae extremely Achromadora species are confined with terrestrial long. Marine. (soil, moss) and, to a lesser degree, freshwater biotopes. Type and only species: Trichethmolaimus hirsutus (Gerlach, 1956) Platt, 1982 ( = Spiliphera hirsuta Gerlach, 1956) 7.12.4 Family Ethmolaimidae Filipjev & Schuurmans 7.12.5 Family Neotonchidae Stekhoven, 1941 Wieser & Hopper, 1966

Diagnosis: Chromadorida. Cuticle with transversal rows Diagnosis: Chromadorida. Cuticle annulated and punc- of punctation, lateral differentiation may be developed as tated. Anterior sensilla arranged in three circles, only the

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.5 Family Neotonchidae Wieser & Hopper, 1966 391

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I

Fig. 7.98 : Examples of Ethmolaimidae and Neotonchidae. A, Ethmolaimus pratensis (from Platt, 1982, Fig. 2); B: Paraethmolaimus appendixocaudatus (from Jensen 1994, Fig. 9); C, Trichethmolaimus hirsuta (from Gerlach 1956, Taf. 31, Fig. A – B); D, Comesa corcunda (from Gerlach 1956, Taf. 30, Fig. A – D); E, Filitonchus ewensis (from Platt 1982, Fig. 36); F, Gomphionchus lutosus (from Wieser & Hopper 1966, Fig. 20); G, Gomphionema typicum (from Platt 1982, Fig. 7 C – E); H, Nannolaimus fusus (from Platt, 1982, Fig. 32); I, Neotonchus punctatus (from Wieser & Hopper 1966, Fig. 1 – 2: anterior body and male tail; Platt 1982, Fig. 16: head).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 392 7.12 Order Chromadorida Chitwood, 1933 four cephalic setae are setiform. Amphideal fovea lateral, Type and only species: Gomphionchus lutosus (Wieser & multispiral with more than three turns. Buccal cavity with Hopper, 1966) Platt, 1982 ( = Neotonchus lutosus Wieser & large dorsal tooth, with or without two small subventral Hopper, 1966) teeth. Pharynx with a muscular end bulb. Two opposed testes, rarely single anterior testis. Midventral precloa- cal cup-shaped supplements present. Spicules short and 7.12.5.4 Genus Gomphionema Wieser & fairly straight but bent one-third of the distance from the Hopper, 1966 (Fig. 7.98 G) distal end. Tail conical, with rounded tip. Strictly marine. Type genus: Neotonchus Cobb, 1933 Diagnosis : Neotonchidae. Buccal cavity heavily cuticu- No morphological holapomorphies are known for the larized, consisting of an anterior concave chamber and a Neotonchidae (Lorenzen 1981, 1994). Characterization of cylindroconoid posterior portion, and armed with a massive Neotonchidae presents a mixture of features of the families dorsal tooth not entering the cheilostoma; subventral teeth Chromadoridae (three separate circles of anterior sensilla, absent. Pharyngeal bulb large, “ barrel-shaped ” , occupying cup-shaped precloacal supplements), Cyatholaimidae approximately one-third of the total pharyngeal length. (multispiral amphideal fovea, diorchic condition) and even Number of species: 3 Microlaimidae (buccal cavity of Comesoma), and therefore Type species: Gomphionema typica Wieser & Hopper, 1966 various neotonchid genera were formerly placed in diffe- rent families. Even the taxon Neotonchidae (originally esta- blished as a subfamily) has been referred to different fami- 7.12.5.5 Genus Nannolaimus Cobb, 1933 lies, Cyatholaimidae and Ethmolaimidae in two revisions of (Fig. 7.98 H) Wieser & Hopper (1966) and Platt (1982). Also, discrimina- tion of now established genera is not always clear. Diagnosis: Neotonchidae. Body cylindrical, filiform. Inner and outer labial sensilla setose. Buccal cavity rather narrow; dorsal tooth small, if present. 7.12.5.1 Genus Comesa Gerlach, 1956 Number of species: 4 (Fig. 7.98 D) Type species: Nannolaimus guttatus Cobb, 1933

( = Neotonchoides Platt, 1982) Diagnosis : Neotonchidae. Buccal cavity with a distinct 7.12.5.6 Genus Neotonchus Cobb, 1933 forward-pointing tooth and ventral ridges but no subven- (Fig. 7.98 I) tral teeth. Posterior pharyngeal bulb small ( < 25% total pharynx length) but distinct. ( = Heterocyatholaimus Allg é n, 1935) Number of species: 9 Diagnosis: Neotonchidae. Body punctated cuticle usually Type species: Comesa corcunda Gerlach, 1956 with some lateral differentiation. Sublateral pores asso- ciated with little somatic setae present. Buccal cavity 7.12.5.2 Genus Filitonchus Platt, 1982 weakly to moderately cuticularized, armed with a small to medium-sized triangular dorsal tooth, and with or (Fig. 7.98 E) without subventral teeth. Pharyngeal end bulb small, dis- tinct. Gubernaculum simple, rod-like. Diagnosis: Neotonchidae. Body cylindrical, filiform. Inner Number of species: 6 labial sensilla papillose whereas outer labial and cephalic Type species: N. punctatus Cobb, 1933 sensilla setose. Buccal cavity rather narrow; dorsal tooth small, if present. Type and only species: Filitonchus filiformis (Warwick, 7.12.6 Family Selachinematidae 1971) Platt, 1982 ( = Neotonchus filiformis Warwick, 1971) Cobb, 1915 7.12.5.3 Genus Gomphionchus Platt, 1982 ( = Choanolaimidae De Coninck & Stekhoven, 1933) (Fig. 7.98 F) Diagnosis : Chromadorida. Annulated cuticle with trans- verse rows of dots often focused as tiny asterisks. Pattern Diagnosis: Neotonchidae. Similar to Gomphionema but of anterior sensilla six + ten. Amphideal fovea lateral, the dorsal tooth less strongly cuticularized and reaching multispiral. Pharynx wide and strongly muscular, often the cheilostoma. with anterior and posterior muscular thickenings. Buccal

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.6 Family Selachinematidae Cobb, 1915 393 cavity well-developed, two-chambered. Buccal armament 7.12.6.1 Subfamily Choniolaiminae derived from the sclerotized structures of the stoma walls Schuurmans-Stekhoven & (rhabdions), in the form of either three sets of small teeth, Adam, 1931 denticles, or two or three mandibles, but never dorsal and subventral teeth. Intestine consists of large bulging Diagnosis : Choniolaimidae. Radially symmetrical stoma cells with distinct boundaries. Precloacal supplementary consists of two compartments: an anterior broad, cup- organs, when present, cup-shaped or setose but never shaped chamber and a posterior narrow, cylindroid tubular. Testes usually paired. Marine. chamber. The anterior chamber is reinforced by six radial Type genus: Selachinema Cobb, 1915 cuticularized rhabdions that may bear tiny denticles pos- Broad truncated cephalic end and voluminous teriorly. Walls of the posterior chamber consist of three complex buccal cavity with elaborated armament are weaker longitudinal rhabdions. developed in connection with the raptorial mode of life Type genus: Choniolaimus Ditlevsen, 1918 of the selachinematids. Many of them were recorded repeatedly as predators of other nematodes. Gerlach (1964a) derived radially-symmetrical, two-chambered 7.12.6.1.1 Genus Choanolaimus de Man, 1880 stoma of selachinematids from the bilaterally-symme- (Fig. 7.99 A) trical stoma of Cyatholaimidae through reduction of the anteriormost vestibular chamber (cheilostoma), Diagnosis : Choniolaiminae. All the anterior sensilla are disappearance of the prominent dorsal tooth and deve- papilloid. Cuticle is laterally differentiated with larger lopment of movable two-chambered stoma with walls more widely separated dots. Amphideal fovea coiled in strengthened by radial sclerotized beams (rhabdions). two or three turns. Rhabdions of the anterior cup-shaped There are two different trends of stoma development in stoma chamber posteriorly bicuspid. Pharynx cylindrical. two subfamilies. In Choniolaiminae, tooth-like struc- Precloacal supplements are papilla pierced by fine pores. tures are differentiated at the posterior ends of the Tail very short, rounded conical or semicircular. rhabdions of the anterior cup-shaped stoma chamber. Type and only species: Choanolaimus psammophilus de When the mouth opens by gripping a prey, the ante- Man, 1880. The species lives mainly in brackish subsoil rior chamber widens funnel-like. Anterior rhabdions water and thus cannot be considered as true marine. diverge distally and meet together proximally, thus holding the prey with the teeth. In contrast, in Sela- chinematinae, when the thin-walled anterior chamber 7.12.6.1.2 Genus Choniolaimus Ditlevsen, 1918 widens, three or two strong indented mandibles of the (Fig. 7.99 B) posterior chamber move outside and open to grasp a prey. There is a sequence of graduated reduction of the ( = Bulbopharyngiella Allg é n, 1929) dorsal mandible within Selachinematinae, from Syn- Diagnosis : Choniolaiminae. Outer labial and cephalic onchium and Synonchiella with three equal mandibles, sensilla setose. Lateral differentiation of cuticle absent. through Kosswigonema in which the dorsal mandible Amphideal fovea coiled in many turns. Anterior cup- is notably smaller than subventral ones, to Demonema shaped stoma chamber relatively small, and its rhabdi- and especially Cheironchus, in which the dorsal mandi- ons only slightly movable; posterior cylindrical stoma ble is vestigial. chamber with strongly sclerotized rhabdions pointed Feeding of selachinematids on other nematodes was anteriorly. Rhabdions of the anterior cup-shaped stoma established already by pioneering nematologists (Allgé n chamber posteriorly bicuspid. Pharynx thickened anteri- 1939, and sources cited therein). Okhlopkov (2003) orly and terminated with a posterior muscular bulb. studied gut content in some selachinematid species in Number of species: 5 the White Sea, Northern Russia and found that Gamma- Type species: Choniolaimus papillatus Ditlevsen, 1918 nema rapax and Halichoanolaimus robustus , juveniles and adults, feed on only nematodes of various families and sizes. On the contrary, the gut content of Latronema 7.12.6.1.3 Genus Cobbionema Filipjev, 1922 aberrans consisted essentially of particles and debris of (Fig. 7.99 C) unclear origin. There are two subfamilies classified within the family Diagnosis: Choniolaiminae. Outer labial sensilla as long Selachinematidae. setae, cephalic sensilla as short setae. Amphideal fovea

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A B

C

D E F

H G I

K J

Fig. 7.99 : Images of genera Selachinematidae. A, Choanolaimus psammophilus (from Platt & Warwick 1988, Fig. 132); B, Choniolaimus panicus (from Okhlopkov 2002, Fig. 1); C, Cobbionema acrocerca (from Stekhoven 1950, Fig. 60); D, Gammanema rapax (from Okhlopkov, 2002, Fig. 6); E, Halichoanolaimus sp. (orig., Okhlopkov); F, Latronema aberrans (from Okhlopkov 2002, Fig. 9– 10); G, Cheironchus conicaudatus (from Tchesunov & Okhlopkov 2006, Fig. 4 – 5); H, Demonema rapax (from Gerlach 1958, Fig. 1 B– C); I, Kosswigonema acanthum (from Gerlach 1964, Abb. 15); J, Synonchiella hopperi (from Tchesunov & Okhlopkov 2006, Fig. 10– 11); K, Synonchium obtusum (from Gerlach 1964, Abb. 16).

Authenticated | [email protected] Download Date | 2/15/14 10:06 AM 7.12.6 Family Selachinematidae Cobb, 1915 395 spirally coiled in three turns. Stoma consists of anterior but distinct annulations and 12– 50 longitudinal ridges. cup-shaped and posterior cylindrical chambers with All anterior sensilla setose, arranged in two circles, six denticles between them. Apophyses projected from ante- + ten; outer labial and cephalic setae not united in late- rior rhabdions into the pharyngeal tissue. Pharynx with romedian pairs but widely separated. Amphideal fovea strong anterior bulb and weaker posterior thickening. Tail often rather small, round, transversally oval, spirally consists of anterior conical and posterior slender cylindri- coiled in one or two turns. Anterior stoma chamber large cal portions. and can open up very wide. Anterior rhabdions very Number of species: 2 strong and terminate posteriorly in many pointed teeth Type species: Cobbionema acrocerca Filipjev, 1922 that protrude outward by the wide opening of the mouth. Supplementary organs sucker-like or cup-shaped. Tail conical. 7.12.6.1.4 Genus Gammanema Cobb, 1920 Number of species: 8 (Fig. 7.99 D) Type species: Latronema orcinum (Gerlach, 1952) Wieser, 1954 ( = Synonchiella orcina Gerlach, 1952) ( = Trogolaimus Cobb, 1920, opinion of Tchesunov & Okhlopkov 2006) Diagnosis : Choniolaiminae. No lateral differentiation of 7.12.6.2 Subfamily Selachinematinae the cuticle. Anterior and cephalic sensilla setose. Four Cobb, 1915 cervical setae developed in most species. Cuticular rhab- dions of the anterior cup-shaped stoma chamber often Diagnosis : Selachinematidae. Anterior chamber of the with fine longitudinal striation; the anterior rhabdi- stoma is reduced. Three rhabdions of the posterior stoma ons posteriorly terminate in teeth-like projections but chamber are modified into powerful, denticulate protrusi- no minute denticles on bottom of the anterior stoma ble mandibles. chamber. No posterior pharyngeal bulb. Tail conical. Type genus: Selachinema Cobb, 1915 Preanal midventral supplementary organs usually present, cup-shaped. Number of species: 9 7.12.6.2.1 Genus Cheironchus Cobb, 1917 (Fig. 7.99 G) Type species: Gammanema ferox Cobb, 1920 ( = Dignathonema Filipjev, 1918) Diagnosis: Selachinematinae. No lateral differentiation in 7.12.6.1.5 Genus Halichoanolaimus De Man, 1886 the cuticle. Short somatic setae or papillae and irregular (Fig. 7.99 E) pores along the body. Three mandibles: two large latero- ventral mandibles, equal in shape and size and shifted ( = Smalsundia Allg é n, 1929) laterally, whereas dorsal mandible is vestigial. Pharynx Diagnosis : Choniolaiminae. Lateral dots of the cuticle are with anterior and posterior muscular thickenings. Supple- bigger and more widely separated. All anterior sensilla are mentary organs cup-shaped. papilloid. Cervical setae not evident. There are three trans- Number of species: 7 versal sets of denticles between the anterior and posterior Type species: Cheironchus vorax Cobb, 1917 stoma chambers. No posterior pharyngeal bulb. Adult stages, at least, are devoid of rectum and anus, i.e., their intestine is blind. Precloacal supplementary organs papilloid or setose. 7.12.6.2.2 Genus Demonema Cobb, 1894 Tail elongated with a distal cylindrical of filiform portion. (Fig. 7.99 H) Number of species: 23 Type species: Halichoanolaimus robustus (Bastian, 1865) ( = Selachinema Cobb, 1915) ( = Spiliphera robusta Bastian, 1865) Diagnosis : Selachinematinae. All the anterior sensilla papilloid. Three mandibles: two large equal lateroven- tral mandibles distally knobbed and consisting of two 7.12.6.1.6 Genus Latronema Wieser, 1954 (Fig. 7.99 F) parts, whereas the dorsal mandible is vestigial. Pharynx cylindrical. Supplementary organs small, setose. Tail Diagnosis : Choniolaiminae. Body very short, cylindrical, consists of anterior conical and posterior long cylindri- with abruptly truncated anterior end. Cuticle with fine cal portions.

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Number of species: 2 Literature Type species: Demonema rapax Cobb, 1894 Allg é n, C. A. (1939): Rä uberische Ernä hrungsweise mariner Nematoden, insbesondere Halichoanolaimini. Folia Zool. Hydrobiol. 9: 321 – 325. Kosswigonema 7.12.6.2.3 Genus Gerlach, 1964 Belogurov, O. I., Dashchenko, O. I. & Fadeeva, N. P. (1985): A (Fig. 7.99 I) free-living nematode Panduripharynx pacifica sp. nov. (Nematoda, Chromadoridae) from the Japan Sea. Nauchnye Diagnosis : Selachinematinae. Four long lateromedian doklady vysshei shkoly. Biol. Nauki 6: 37 – 42. setae and six lateral and submedian pairs of short setae. Boucher, G. (1976): N é matodes des sables fins infralittoraux de la Pierre Noire (Manche occidentale) II. Chromadorida. Bulletin du Amphideal fovea spirally coiled in 2.5– 3 turns. Stoma Mus é um national d ’ histoire naturelle 3e ser., no 352. Zoologie consists of cup-shaped anterior chamber with strongly 245: 25 – 61. sclerotized walls and posterior portion transformed into Boucher, G. & de Bov é e, F. (1971): Trochamus carinatus gen. et sp. n. a mandibular apparatus. Three mandibles: two sub- et Adeuchromadora megamphida gen. et sp. n. Chromadoridae ventral mandibles strong, consisting of two parts and (Nematoda) à dix soies c è phaliques de la vase terrig è ne c ô ti è re bearing numerous teeth, and the dorsal mandible weaker. de Banyuls-sur-Mer. Vie et Milieu 22 A: 231 – 242. Blome, D. (1974): Zur Systematik von Nematoden aus dem Sandstrand Pharynx muscular, anteriorly swollen; no posterior bulb. der Nordseeinsel Sylt. Mikrofauna des Meeresbodens 33: 75– 99. Supplementary organs cup-shaped. Tail short, conical. Blome, D. (1982): Systematik der Nematoda eines Sandstrandes der Type and only species: Kosswigonema acanthum (Gerlach, Nordseeinsel Sylt. Mikrofauna des Meeresbodens 86: 1 – 194. 1957) Gerlach, 1964 ( = Selachinema acanthum Gerlach, Coles, J. W. (1965): A critical review of the marine nematode genus 1957) Euchromadora de Man, 1886. Bull. Br. Mus. Nat. Hist., Zool. 12: 157 – 194. De Coninck, L. A. & Schuurmans Stekhoven, J. H. (1933): The free-living marine nemas of the Belgian coast II. M é moires du 7.12.6.2.4 Genus Synonchiella Cobb, 1933 Mus é e royal d ’ Histoire naturelle de Belgique 58: 1 – 163. (Fig. 7.99 J) De Mesel, I., Lee, H. J., Vanhove, S., Vincx, M. & Vanreusel, A. (2006): Species diversity and distribution within the deep-sea Diagnosis : Selachinematinae. Cuticle with transverse rows nematode genus Acantholaimus on the continental shelf and slope in Antarctica. Polar Biol. 29: 860 – 871. of dots arranged transversally or longitudinally; lateral dif- Dashchenko, O. I. (2002): Three new free-living marine nematode ferentiation of the cuticle weakly developed, lateral diffe- species of the subfamily Hypodontolaiminae (Chromadorida, rentiation or not developed at all. Anterior stoma chamber Chromadoridae). Zoologicheskii Zhurnal 81: 771 – 778 nonsclerotized and inconspicuous, posterior stoma equip- [in Russian, English summary]. ped with three equally solid mandibles distally biliobed, Dashchenko, O. I. & Belogurov, O. I. (1984): The morphology of Ptycholaimellus adocius sp. n. (Nematoda, Chromadorida) with hooks or claws anteriorly. Pharynx without posterior from sponges of the Posjet Bay, Sea of Japan. Zoologicheskii bulb. Cup-shaped precloacal supplementary organs. Tail Zhurnal 63: 976 – 982 [in Russian, English summary]. conical or with hind cylindrical portion. Decraemer, W. & Smol, N. (2006): Orders Chromadorida, Number of species: 15 Desmodorida and Desmoscolecida. In: Eyualem Abebe, A., Type species: Synonchiella truncata Cobb, 1933 Traunspurger, W. & Andr á ssy, I. (eds.) Freshwater Nematodes: Ecology and Taxonomy , pp. 497 – 573. CABI, Wallingford. Eskin, R. A. & Hopper, B. (1985): Population dynamics and description of Ptycholaimellus hibernus n. sp. (Nematoda: 7.12.6.2.5 Genus Synonchium Cobb, 1920 (Fig. 7.99 K) Chromadoridae). J. Nematol. 17: 38 – 45. Filipjev, I. (1917): Un n é matode libre nouveau de la mer Caspienne, Diagnosis : Selachinematinae. Cuticle with transver- Chromadorissa gen. nov. (Chromadoridae, Chromadorini). sal rows of dots. Anterior sensilla papilloid. Amphideal Zoologicheskii Zhurnal 2: 24 – 30. Gagarin, V. G. (2001): Descriptions of free-living nematodes spirally coiled, relatively small and transversally oval. Achromadora sedata sp. n. and A. chungsani (Hoeppli, Chu, Anterior stoma chamber cup-shaped and relatively volu- 1932) (Chromadorida), for the first time discovered at Ukraina. minous, with weak rhabdions; posterior stoma chamber Biologia Vnutrennikh Vod 1: 11 – 17. provided with three equal mandibles with strong median Gerlach, S. A. (1951): Nematoden aus der Familie der Chromadoridae tooth and weaker flank teeth. Preanal supplementary von den deutschen Kü sten. Kieler Meeresforschungen 8: 106 – 132. organs inconspicuous or absent. No gubernaculum. Tail Gerlach, S. A. (1953a): Die Nematodenfauna der Uferzonen und des K ü stengrundwassers am finnischen Meeresbusen. Acta Zool. short, conical. Fenn. 73: 1 – 32. Number of species: 3 Gerlach, S. A. (1953b): Recherches sur la faune des eaux interstitielles Type species: Synonchium obtusum Cobb, 1920 de Madagascar. III. Sur quelques Né matodes libres des eaux

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souterraines littorales de Madagascar. M é moires de l’ Institut Jensen, P. (1980): Description of the marine free-living nematode scientifique de Madahgascar (A) 8: 73 – 86. Chromadora lorenzeni n. sp. with notes on its microhabitats. Gerlach, S. A. (1953c): Die Nematodenbesiedlung des Sandstrandes Zoologischer Anzeiger (Jena) 205: 213 – 218. und des K üstengrundwassers an der italienischen K ü ste. I. Jensen, P. (1994): Revision of Ethmolaiminae Filipjev & Stekhoven, Systematischer Teil. Archivio Zoologico Italiano 37: 517 – 640. 1941 (Nematoda, Chromadorida), with descriptions of one new Gerlach, S. A. (1956): Diagnosen neuer Nematoden aus der Kieler genus and three new species. Hydrobiologia 286: 1 – 15. Bucht. Kieler Meeresforschungen 12: 85 – 109. Kito, K. (1978): Five species of marine nematodes of the genus Gerlach, S. A. (1958): Freilebende Nematoden von den Korallenriffen Chromadora Bastian from Japan. Annotationes Zoologicae des Roten Meeres. Kieler Meeresforschungen 14: 241 – 246. Japonensis 51: 164 – 178. Gerlach, S. A. (1964a): Revision der Choniolaiminae und Kito, K. (1977): Studies of the free-living marine nematodes from Selachinematinae (freilebende Meeres-Nematoden). Hokkaido, II. Proc. Jpn. Soc. Syst. Zool . 13: 17 – 23. Mitteilungen aus dem Hamburgischen Zoologischen Lorenzen, S. (1972a): Die Nematodenfauna im Verklappungsgebiet fü r Museum Kosswig-Festschrift 23 – 50. lndustrieabwasser nordwestlich von Helgoland. II. Desmodorida Gerlach, S. A. (1964): Neue Cyatholaimidae (Nematoda Chromadorida) und Chromadorida. Zoologischer Anzeiger 187: 283– 302. von den Malediven. Ver ö ffentlichungen des Instituts f ü r Lorenzen, S. (1972b): Die Nematodenfauna im Verklappungsgebiet Meeresforschungen in Bremerhaven 9: 70 – 78. f ü r Industrieabw ä sser nordwestlich von Helgoland. III. Gerlach, S. A. (1965): Freilebende Meeresnematoden aus der Cyatholaimidae, mit einer Revision von Pomponema Cobb 1917. Gezeitenzone von Spitzbergen . Verö ffentlichungen des Ver ö ffentlichungen des Instituts f ü r Meeresforschungen in Instituts fü r Meeresforschungen in Bremerhaven 9: 109 – 172. Bremerhaven 13: 285 – 306. Gerlach, S. A., Schrage, M. & Riemann, F. (1979): Die Gattung Lorenzen, S. (1981): Entwurf eines phylogenetischen Systems der Acantholaimus (Nematoda, Chromadoridae), und Beobachtungen freilebenden Nematoden. Ver ö ffentlichungen des Instituts f ü r ü ber einen mutmasslichen Transportmechanismus f ü r Spermien Meeresforschung in Brem e rhaven 7, Suppl.: 472. bei A. calathus sp. n. Ver ö ffentlichungen des Instituts fü r Lorenzen, S. (1994): The Phylogenetic Systematic of Freeliving Meeresforschungen in Bremerhaven 18: 35– 67. Nematodes . The Ray Society, London. Gal ’ tsova, V. V. (1985): Free-living marine nematodes as a component Miljutina, M. A., Miljutin, D. M. & Tchesunov, A. V. (2013): Seven of the meiobenthos of Chupa Inlet of the White Sea. In: Acantholaimus (Chromadoridae: Nematoda) species from one Platonova T. A. & Gal’ tsova, V. V. (eds.) Nematodes and Their Role deep-sea sediment sample (Angola Basin, south-east Atlantic). in the Meiobenthos , pp. 165 – 270. Nauka, Leningrad [Translation J. Mar. Biol. Assoc. UK 93: 935 – 953. from Russian, published for the Smithsonian Institution Miljutina, M. A. & Miljutin, D. M. (2012): Seven new and four Libraries and the National Science Foundation, Washington, D.C. known species of the genus Acantholaimus (Nematoda: by Amerind Publishing Co. Pvt. Ltd., New Delhi]. Chromadoridae) from the abyssal manganese nodule field Holterman, M., Holovachov, O., van den Elsen, S., van Megen, (Clarion-Clipperton Fracture Zone, North-Eastern Tropical H., Bongers, T., Bakker, J. & Helder, J. (2008): Small subunit Pacific). Helgol. Mar. Res . 66: 413 – 462. ribosomal DNA-based phylogeny of basal Chromadoria Murphy, D. G. (1963): A new genus and two new species of (Nematoda) suggests that transitions from marine to terrestrial nematodes from Newport, Oregon. Proc. Helminthol. Soc. habitats (and vice versa ) require relatively simple adaptations. Washington 30: 73 – 78. Mol. Phylogenet. Evol. 48: 758 – 763. Murphy, D. G. (1965): The marine nematode genus Nygmatonchus Hope, W. D. & Murphy, D. G. (1972): A taxonomic hierarchy and Cobb, 1933 rediscovered, with description of N. alii , new checklist of the genera and higher taxa of marine nematodes. species. Verö ffentlichungen des Instituts fü r Meeresforschung Smithsonian Contributions to Zoology 137: 1 – 101. in Bremerhaven 9: 204 – 209. Hopper, B. E. (1961): Marine nematodes from the coast line of the Okhlopkov, J. R. (2002): Free-living nematodes of the families Gulf of Mexico II. Can. J. Zool. 39: 359 – 365. Selachinematidae and Richtersiidae in the White Sea Hopper, B. E. (1963): Punctodora exochopora n. sp. from the (Nematoda, Chromadoridae). Zoosyst. Rossica 11: 41 – 55. Canadian shore of Lake Ontario. Can. J. Zool. 41: 1121 – 1126. Okhlopkov, J. R. (2003): Nutrition of free-living nematodes of the Hopper, B. E. (1966): Theristus polychaetophilus n. sp. (Nematoda), families Selachinematidae and Richtersiidae in the White Sea. an external parasite of the spionid polychaete Scolelepis Proceedings of the Pertsov White Sea Biological Station, ( Scolelepis) squamata Mü ller, 1806. Can. J. Zool. 44: 787 – 791. Vol. 9. Proceedings of the 7 th International Conference , Hopper, B. E. (1972): Free-living marine nematodes from Biscayne p. 127 – 139, 10 – 11. August, 2002. KMK Scientific Press, Bay, Florida IV. Cyatholaimidae: on the occurence of Marylina Moscow. [in Russian]. n. gen. and Longicyatholaimus Micoletzky, 1924 in Biscayne Ott, J. A. (1972): Twelve new species of nematodes from an intertidal Bay, with a description of L. longicaudatus (De Man, 1876) from sandflat in North Carolina. Internationale Revue der gesamten the type locality. Zoologischer Anzeiger (Jena) 189: 64 – 88. Hydrobiologie 57: 463 – 496. Huang, Y. & Zhang, Z. (2007): A new genus and new species of Pastor de Ward, C. T. (1985): Free-living marine nematodes of the free-living marine nematodes from the Yellow Sea, China. Deseado river estuary (Chromadoroidea: Chromadoridae, J. Mar. Biol. Assoc. UK 87: 717 – 722. Ethmolaimidae, Cyatholaimidae and Choniolaimidae) Santa Inglis, W. G. (1963): New marine nematodes from off the coast of Cruz, Argentina. 5. Publicaci ó n Especial del Centro Nacional South Africa. Bull. Brit. Mus. Nat. Hist. 10: 529 – 552. Patag ó nico 6: 1 – 83. Inglis, W. G. (1969): Convergence in the structure of the head Platonova, T. A. & Potin, V. V. (1972): On new genera Harpagonchus and cuticle of Euchromadora species and apparently similar and Harpagonchoides (Nematoda, Chromadorida, nematodes. Bull. Brit. Mus. Nat. Hist., Zool. 17: 149 – 204. Harpagonchidae fam. n.) living on the parapodia and gills of

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the antarctic polychates Aglaophamus Kinberg and Hemipodus Timm, R. W. (1969): Megodontolaimus new genus (Nematoda: Quatrefages. Issledovanija sovjetskichn antarktitcheskich Chromadoridae), with description of two new species. Proc. Ekspedizii 5: 81 – 85 [in Russian, English summary]. Helminthol. Soc. Washington 36: 36 – 39. Platt, H. M. (1982): Revision of the Ethmolaimidae (Nematoda: Vitiello, P. (1970): N é matodes libres marins des vases profondes du Chromadorida). Bull. Brit. Mus. Nat. Hist., Zool. 43: 185 – 252. Golfe du Lion. II. Chromadorida. T é thys 2: 449 – 500. Platt, H. M. & Warwick, R. M. (1988): Free-Living Marine Nematodes. Warwick, R. M. & Coles, J. W. (1975): Notes on the free-living Part II. British Chromadorids. Pictorial Keys to World Genera marine genus Euchromadora de Man, 1886 and its allies, with and Notes for the Identification of British Species . Synopses of descriptions of two new species (Chromadoridae: Nematoda). the British Fauna (Nerw Series), 38. Published for the Linnean J. Nat. Hist. 9: 403 – 412. Society of London and the Estuarine and Brackish-Water Wieser, W. (1954): Free-living marine nematodes. II. Chromadoroidea. Sciences Association. By E. G. Brill / Dr. W. Backhuys, Leiden, Lunds Universitets Å rsskrift. N. F. Avd. 2 50: 1– 148. New York, K Ø benhavn, K ö ln. Wieser, W. (1955): Eine Sammlung mariner Nematoden aus Piraeus Platt, H. M. & Zhang, Z. N. (1982): New species of marine nematodes (Griechenland). Ö sterreichische zoologische Zeitschrift 6: from Loch Ewe, Scotland. Bull. Brit. Mus. Nat. Hist., Zool. 42: 597 – 630. 227 – 246. Wieser, W. (1959): Free-Living Nematodes and Other Small Sharma, J. & Vincx, M. (1982): Cyatholaimidae (Nematoda) from the Invertebrates of Puget Sound Beaches . University of Canadian Pacific coast. Can. J. Zool. 60: 271 – 280. Washington Press. Seattle. Tchesunov, A. V. (2008): Three new species of free-living nematodes Wieser, W. (1968): Chromadorina astacicola (Schneider, 1932) from the South-East Atlantic abyss (DIVA I Expedition). Zootaxa und Chromadorina majae n. sp., zwei mit Decapoden 1866: 151 – 174. vergesellschaftete Nematoden. Thalassia Jugoslavica 4: 39– 43. Tchesunov, A. V. & Okhlopkov, Yu. R. (2006): On some selachinematid Wieser, W. & Hopper, B. (1966): The Neotonchinae, new subfamily nematodes (Chromadorida: Selachinematidae) deposited in (Cyatholaimidae: Nematoda), with an analysis of its genera, the collection of the Smithsonian National Museum of Natural Neotonchus Cobb, 1933 and Gomphionema new genus. Can. History. Nematology 8: 21– 44. J. Zool. 44: 519 – 532. Timm, R. W. (1961): The marine nematodes of the Bay of Bengal. Wieser, W. & Hopper, B. (1967): Marine nematodes of the east coast of Proc. Pak. Acad. Sci. 1: 1 – 88. North America. I. Florida. Bull. Mus. Comp. Zool. 135: 239– 344.

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