The freeliving marine nematode genus

Transcription

1 <oulogical Journal of Ihe Linnean SocieQ (1985), 83: With 23 figures The freeliving marine nematode genus Sabatieriu (Nematoda: Comesomatidae). Taxonomic revision and pictorial keys HOWARD M. PLATT, F.L.S. Department of,zoology, British Museum (Natural History), Cromwell Road, London S W7 5BD Received October 1983, accepted for publication February 1984 Relationships within the freeliving marine nematode family Comesomatidae and the systematic position of the Comesomatidae are discussed. Composite pictures representing all valid comesomatid genera are provided as an aid to identification. Of the 73 nominal Sabafieria species extant prior to this revision: six were already considered species dubiae, to which is added a further 14 based on insufficient original description; one is transferred to another genus; nine are synonymized with previously described species; seven are considered species inquirendae, to which is added a new species previously described under another name; two possess synapomorphic features warranting recognition as a separate genus, SetosabutirriP gen. nov.; 34 are considered valid Sabatieria species and two previously synonymized species are reinstated. Pictorial keys and tabular information are provided to facilitate species identification. Several other minor taxonomic changes are proposed. KEY WORDS:-Nematoda - Comesomatidae - Sabafieria - marine - taxonomy - revision - pictorial keys. CONTENTS Introduction Materials and methods Systematic position of the Comesomatidae Outline classification of the Comesomatidae Morphology and character evaluation of Sabatieria species 'laxonomic revision and identification Family Comesomatidae Filipjev, Subfamily Sabatieriinae Filipjev, Setosabatieriu gen. nov Sabafieriu Rouville, Rrfercnces Appendix List of valid Sabatieria species List of species inquirendae List of dubious species List of new synonyms INTRODUCTION Species of the genus Sabahria Rouville, 1903 are among the most common freeliving nematodes in marine benthic habitats, especially from mud and muddy sand. The genus is one of the two most frequently reported from British /85/ $03.00/ The Linnean Society of London

2 28 H. M. PLATT coastal waters and Sabatieria species are often the most abundant organisms in nematode assemblages. Over 80 species have been described, although several have subsequently been synonymized or considered dubious. Attention has recently been drawn to the taxonomy and ecology of this important genus by Jensen (1979a, 1981), who provided an update of Wieser s (1954) verbal key. Work involved in the description of new species from Antarctica (Platt, 1983) and the re-examination of some European species (Platt, 1984) highlighted the difficulties involved in separating species of the genus. Consequently a more detailed revision of the whole genus was undertaken. Verbal keys proved unsuitable for such a large number of species so pictorial keys were devised. It also seemed appropriate to attempt to set the genus in context with the other genera of the family Comesomatidae and to comment on the position of the family itself. MATERIALS AND METHODS The methodology and abbreviations used here are as previously described elsewhere (Platt, 1982). In addition to the specimens listed in the species revision section of this paper and in Platt (1982, 1983, 1984), additional outgroup material (being those examples referred to by species name and registration number in the text) were also examined: all the material is in the collection of the British Museum (Natural History). The terms species inquirenda and species dubia are often used in the same sense. However, throughout this work the terms are kept as distinct categories. Species inquirenda is used for those species deemed to be in the appropriate genus but for which there is inadequate information. If specimens of a species inquirenda were ever found again, they would probably be recognized as such and the missing information supplied. Therefore, species inquirendae cannot be ignored. Species dubia, on the other hand, is reserved for those species whose descriptions are so insufficient or generally poor that if specimens were ever found again conspecificity could not be proved beyond any reasonable doubt. In many cases, descriptions based only on juveniles or females render the species dubious because almost invariably the diagnostic characters are vested in the male. For all practical purposes, species dubiae can be ignored. SYSTEMATIC POSITION OF THE COMESOMATIDAE The systematic position of this family is somewhat problematical. According to some authors (e.g. Wieser, 1954; de Coninck, 1965; Gerlach & Riemann, 1973) it is part of the Chromadorida because of the punctated cuticle and spiral amphids. However, others (e.g. Filipjev, 1934; Lorenzen, 1981) include it with the Monhysterida, based on the shared occurrence of outstretched ovaries. Which of these two hypotheses is followed has a profound influence on subsequent decisions of holophyly and is therefore worthy of consideration and discussion at the outset. ( 1) Comesomatidae as Monhysterida. This hypothesis places maximum weight on the structure of the ovaries. However, outstretched ovaries do occur elsewhere in

3 REVISION OF SABATIERIA 29 the Nematoda (Lorenzen, 1981: 112) and are therefore not confined to the Monhysterida/Comesomatidae and in at least one comesomatid genus, Hopperia, reflexed ovaries are known to occur (Jensen, 1979a). Like the comesomatids, many monhysterids also have paired dorso-caudally directed gubernacular apophyses, structures otherwise absent in most families of the Chromadorida although present in other nematodes (e.g. Pseudocella caecum BMNH: , an enoplid). Within the Monhysterida, Lorenzen (1981) also draws attention to the presence of 3 or 6 tooth-like structures in the anterior part of the buccal cavity in some comesomatids (e.g. Paracomesoma, Hopperia) and marginal pharyngeal tubes as possibly similar to the 6 odontia and pharyngeal tubes in some genera of the monhysterid family Axonolaimidae. He groups both the Comesomatidae and Axonolaimidae together with the Diplopeltidae and Coninckidae in the superfamily Axonolaimoidea. However, marginal tubes are found throughout the Nematoda (e.g. Rhabditis, Paracanthonchus). A consequence of this hypothesis is that the transversely punctated cuticle, multi-spiral amphids and precloacal file of tubular supplementary organs are either plesiomorphic for the Monhysterida + Chromadorida (unlikely on grounds of commonality) or developed independently in both orders. (2) Comesornatidae as Chromadorida. This hypothesis places importance on the presence of a transversely punctated cuticle, multi-spiral amphids and precloacal file of tubular supplements. It should be noted, however, that precloacal supplements and punctated cuticle are not present in all species of the Comesomatidae. For example, a punctated cuticle is absent in species of the genera Laimella, Cervonema and Setosabatieria gen. nov. and some species of Paracomesoma, and no Pierrickia or Metacomesoma species have been described with supplements. A consequence of this hypothesis is that outstretched ovaries are either plesiomorphic for Monhysterida + Chromadorida (considered unlikely on grounds of commonality by Lorenzen, 1981) or developed independently at least twice. Also, apophyses on the gubernaculum developed independently in Comesomatidae, Chromadorida and Enoplia and the presence of a transversely punctated cuticle, multi-spiral amphids and a file of precloacal supplements are plesiomorphic for the Comesomatidae. Discussion: Clearly, the resolution of which group the comesomatids should most logically belong to is not simple. Without good morphological evidence to support suspected homologies, such as might be forthcoming from ultrastructural investigations, all we can do is to search for the most parsimonious solution pro tempore. Evidence from pharyngeal marginal tubes, tooth-like structures in the anterior part of the buccal cavity, and paired gubernacular apophyses is equivocal and could be made to support either hypothesis. However, what appears to be significant to me is that within the Chromadorida (excluding comesomatids for the sake of argument) members of the superfamily Chromadoroidea all have transversely punctated cuticles and that the Cyatholaimidae also have the same combination of cuticle + amphid + precloacal supplements as the Comesomatidae. To have arrived at this latter combination of complex characters independently appears to be less probable than that the intrinsically more simple character of outstretched ovaries evolved independently. On this basis, I will follow hypothesis (2). According to the principle of commonality, those few cases within the Comesomatidae where the

4 30 H. M. PLATT cuticle and precloacal supplement characters are absent must be considered apomorphic, i.e. secondarily lost. OUTLINE CLASSIFICATION OF THE COMESOMATIDAE In order to put into context the discussion of important Sabatieria characters, an outline of the relevant differentiating features of all the comesomatid genera seems appropriate. Further detailed discussion can be found in Jensen (1979a). Pictorial keys (Figs 1-3) for each of the three subfamilies (Sabatieriinae, Comesomatinae and Dorylaimopsinae) are also provided as an aid to identification. Generic characters discussed are numbered ( 1-14) to correlate with the pictorial keys. At this point, a strict phylogenetic analysis of the relationships within the Comesomatidae will not be attempted. Family Comesomatidae The characteristic feature shared by all known comesomatid species is outstretched ovaries (except one species of Hopperia). They also have typically a cephalic sensilla pattern, multi-spiral amphids, punctated cuticles, paired apophyses on the gubernaculum, tubular precloacal supplements and conico-cylindrical tails: all these characters may also be found in other nematode taxa. Sub family Sabatieriinae (Fig. 1) I could not find any features which all genera of the subfamily share uniquely so that, for the present, genera of the subfamily can only be described as lacking the derived features of the other two families. Sabatieria Like the subfamily, Sabatieria can only be recognized by lacking the diagnostic features of the other genera. In other words, Sabatieria represents the seminal form from which all related species-groups can be derived. This can be supported by the much larger number of valid species (36) ascribed to Sabatieria compared with any other comesomatid genus, if commonality is accepted as an indication of primitiveness (Arnold, 1981). Sabatieria will be discussed in more detail below. Scho Lpaniella This monotypic genus, based on a single male specimen, is similar to Sabatieria but was described as lacking the typical dorso-caudally directed gubernacular apophyses (l), although some form of apophysis was depicted (Fig. 1). Jensen (1979a) stated that there is a single apophysis but this does not seem to be exactly what Sergeeva (1972) meant: whether the depicted process is single or double is not clear from Sergeeva s description. In any case, the form of the gubernaculum appears unique and can be considered as an autapomorphy. Sergeeva (1972) also considered the tail to be unique but this is not so: cf. Sabatieria lyonessa (Fig. 13).

5 REVISION OF SABATIERIA 3 1 A B C D I E F G 516 2,5,7 5,8 Figure I. Composite pictures representing genera of the subfamily Sabatieriinae. A, Sabatimia; B, Scholpaniella; C, Pierrickia; D, Acfarjania; E, Liamclla; F, Cervonema; G, Setosabafieria. Numbers refer to the following characters: 1, dorso-caudally directed gubernacular apophyses absent; 2, R2 and R3 sensilla of equal length; 3, precloacal supplements absent; 4, longitudinal rows of lateral punctations; 5, cuticle punctations absent; 6, R2 sensilla situated close to R3 sensilla; 7, elongated head region (between R3 sensilla and amphid); 8, sublateral rows of long cervical setae. Pierrickia The three species were described as having the R2 and R3 sensilla of equal length (2) and without supplements (3), the former feature shared by Cervonema and the latter by Metacomesoma. Actarjania This genus is characterized by longitudinal rows of lateral punctations (4). Jensen ( 1979a) synonymized the three species with Sabatieria, commenting, this kind of differentiation is also found in some species of Sabatieria. However, I cannot discover the species to which Jensen referred and so here retain the distinction. Dorylaimopsis also has similar punctations.

6 32 H. M. PLATT Laimella The four species lack cuticle punctations (5) and have the R3 sensilla situated close to the R2 sensilla (6). The former feature also occurs in Cervonema and Setosabatieria gen. nov., the latter in Metacomesoma. Re-examination of the specimens described by Ward (1974) as Laimella jlicaudata BMNH: (subsequently synonymized with L. longicauda Cobb by Jensen, 1979a) failed to reveal the fine punctations referred to in the original description. Cervonema Apart from characters (2) and (5), the three species of this genus have an elongated head (7), a feature not found in any other comesomatid group. Setosabatiericr gen. nov. The two species, discussed further below, lack cuticle punctations (5) and have sublateral rows of long cervical setae (8). Subfarnib Comesomatinae (Fig. 2) Until recently, the unique characters shared by genera of this subfamily were the presence of elongated spicules (9) and the absence of apophyses on the gubernaculum ( 1). However, Gourbault ( 1980) recently described a Comesomalike species (i.e. with long spicules) which had apophyses: she placed it in a new genus, Comesomoides. Whether Comesomoides should be considered as a member of the subfamily Sabatieriinae with elongate spicules or a member of the subfamily Comesomatinae with apophyses is unclear. Gourbault ( 1980) also draws attention to certain similarities with the subfamily Dorylaimopsinae. Pending a more comprehensive analysis of the whole family, it seems simplest to leave Comesomoides in the subfamily to which Gourbault ( 1980) originally assigned the 9 9, I, 10 9, I, II 9,1,3,6 Figure 2. Composite pictures representing genera of the subfamily Comesornatinae. A, Comcsomoides; B, Comesoma; C, Paracomesoma; D, Metacomesoma. Numbers refer to the following characters: 1, 3 and 6, see Fig. 1; 9, spicules elongate; 10, 4 subcephalic sensilla situated close to R3 sensilla; 11, toothlike projections in the anterior part of buccal cavity.

7 REVISION OF SABATIERIA genus, namely the Comesomatinae. As a consequence, only the presence of elongated spicules is now shared by all genera of the Comesomatinae. Comesornoides See above. Monotypic. Comesoma An addition to character (9), the 10 species of this genus lack gubernacular apophyses (1) and also have 4 subcephalic sensilla situated close behind the R3 sensilla ( 10). Paracomesoma In addition to characters (1) and (9), this genus has teeth in the anterior of the buccal cavity (11) and four of the seven species lack cuticle punctations. Metacomesoma In addition to characters (1) and (9), the only valid species lacks precloacal supplements (3) and has the R2 and R3 sensilla close together (6). Metacomesoma is rather poorly known and the possibility exists that precloacal supplements have been overlooked. 3 :?I Sub fam$y Dorylaimopsinae (Fig. 3) All members of this subfamily have the posterior part of the buccal cavity structually expanded (12). Metasa batieria This monotypic genus lacks the teeth of the other genera. Vasostoma This is another monotypic genus, similar to Metasabatieria except that it has three tooth-like projections in the anterior part of the buccal cavity (11). Paramesonchium The two species of this genus have characters (11) and (12) but differ from all others of the subfamily in having a conical rather than cylindrical buccal cavity (13). Hopperia The only two valid species of Hopperia have characters (1 1) and (12) but are distinguished by the degree of development of the teeth, which are wellcuticularized solid structures (14). In one species, the ovaries were reported to be reflexed. A third species, H. doylaimopsoides (Allgh, 1959) was referred to this genus by Jensen (1979a) but the original description is poor and the species is here considered dubious. Dorylaimopsis The 11 Dorylaimopsis species have characters (11) and (12) but are characterized by having longitudinal files of lateral punctations (4). 3

8 H. M. PLATT C 12 12, II 12, II, 13 D E 12, II, 14 12, 11,4 Figure 3. Composite pictures representing genera of the subfamily Dorylaimopsinae. A, Metasabatieria; B, Vasostoma; C, Paramesonchium; D, Hopperia; E, Dorylaimopsis. Numbers refer to the following characters: 4 and 11, see Figs 1 and 2; 12, posterior part of buccal cavity structurally expanded; 13, buccal cavity conical; 14, teeth solid cuticularized structures. MORPHOLOGY AND CHARACTER EVALUATION OF SABATIERIA SPECIES In the following discussion of the important morphological characters of valid Sabatieria species, an attempt will be made to indicate the taxonomic level of universality at which each appears to be operating. Features of comesomatids in general were discussed above and in detail by Jensen (1979a) and will not be repeated here. During the course of reviewing Sabatieria, the species appeared to fall into five distinct groups, most of which had autapomorphies. These five groups will be referred to by their better known species: praedatrix group, armata group, pulchra group, celtica group and ornata group. Characters of each group are lettered

9 proedutr ix group REVISION OF SABATlEKI,4 urmafo group pulchro group :I celtico group ornofa group \ b, f b, f, 9 Figure 4. Composite pictures representing the five subgroups of the genus Subahria: pruedutris group, arrnutu group, pulchra group, cellicu group, ornnta group. Letters refer to the following characters: a, bilaterally symmetrical elongated cervical setae; b, conspicuous precloacal supplements; c, cuticularized bar on cuneus of gubernaculum; d, ventro-sublateral pair of cervical setae; c, anterior group of more closely situated precloacal supplements; f, guhrrnacular apophyscs curved; g, posterior group of more closely situated precloacal supplements. (a-g) to correlate with the pictorial key (Fig. 4). The species assigned to each group are listed in Tables 2-6. In addition to the five Sabatieria groups, some species were considered to constitute a distinct genus, as described below. Body shape: Most adult Sabatieria species are typical of sediment dwelling nematodes, being pm long with an a -ratio (total body length divided by the maximum body diameter) in the range. Only one species

10 36 H. M. PLATT

11 REVISION OF SABATIERIA :3 7 has so far been measured at over 4000 pm: S. demani at 8000 pm. Species in the pulchra group tend to be the shortest (e.g. S. propissina, pm) while the armata group are the thinnest, with an a -ratio in the range (except S. migrans, see Table 3). There appears not to be a typical fixation-shape, although they are seldom coiled. There is usually a marked degree of anterior attenuation, the head diameter usually being less than half that of the body at the base of the oesophagus. Normally the attenuation is gradual (Fig. 5E) but in one species, S. kelletti (Fig. 5B), the reduction in width occurs conspicuously in the post-amphid region. The tail is always conico-cylindrical (Fig. 51) although in the pulchra group the cylindrical part is usually reduced (Fig. 55). Features of body shape are not unique statements for Sabatieria species and are of little phylogenetic use. Cuticle ornamentation: The annulated cuticle is punctated (Fig. 6). However, S. hilarula was found to be devoid of these dots, only annulations being visible: together with the longitudinal rows of cervical setae, this was considered sufficient to justify placing this and related species into a separate genus. In all cases, the dots are more or less arranged in transverse rows. In the mid-body region of some species (Fig. 6A, B), there the same number of transverse rows on the lateral field as sublaterally, although the dots may appear slightly larger (Fig. 6D). In the anterior oesophageal and caudal regions, there is a more marked tendancy for the lateral dots to be larger than the medial dots, more irregularly arranged (Fig. 6C) and also to be larger than the lateral dots in the mid-body region (Fig. 6C, D). As well as having larger dots in the lateral field, there may also be fewer transverse rows (Fig. 6E, F) which may or may not be confined to the oesophageal and caudal regions. Finally, in some species, the coherence of the transverse rows breaks down in the lateral field (Fig. 6G) so that the dots appear to be randomly distributed. A transversely punctated cuticle is found in other genera of the Comesomatidae as well as in several other chromadorid families, e.g. Cyatholairnidae, Selachinematidae, Ethmolaimidae; as does the lateral differentiation of larger and/or fewer rows of dots. The presence of anteroposterior differentiation of larger dots in the anterior and posterior regions is also a common phenomenon (e.g. Pomponema reductum BMNH: , Paracanthonchus inglisi BMNH: ). Therefore, the variable cuticle condition in Sabatieria must be judged as the discontinuous expression of a complex series of symplesiomorphic features. The exact form can also show a degree of infra-specific variation (cf. S. celtica in Fig. 6E, F). Together with the fact that in many species descriptions too little emphasis has been placed on the antero-posterior differentiation, it means that in separating Sabatieria species no great reliance can be placed on descriptions of the cuticle ornamentation. It also suggests that more attention should in future be paid to cuticle details. Figure 5. Morphological features of Sabafierza species. A, Amphid, S. kelletti; B, anterior oesophageal region, S. kelletti; C, anterior cloacal region, S. kelletti; D, amphid, S. hilarula; E, oesophageal region, S. breuiseta; F, anterior cloacal region, S. breviseta; G, dorsal view of opposed testes of S. ornala; upper arrow indicates start of anterior testis, left of gut and posterior arrow indicates start of posterior testis, right of gut; H, anterior cloacal region,.s. ornoto; I, tail, S. ornnta; J, tail, S. brez~zsefa. Scale bars: A, D = 10 pm; F,I = 30 pm; B, C, E, G, HJ = 50 pm.

12 38 H. M. PLATT Figure 6. Lateral view of cuticle ornamentation. A, S. alata, midbody region; B, S. clongata, midbody region; C, D, S. pulchra from Exe estuary, posterior oesophagus region (C) and mildbody region (D); E, S. celtica from Clew Bay, midbody region; F, S. kelletti, midbody region; G, S. puncfato, posterior oesophagus region. Scale bar = 10 pm.

13 REVISION OF SABATIERIA.j9 Somatic setae: There are usually four sublateral rows of short somatic setae running the length of the body. In some species, the setae in the cervical region may be longer and/or have a specific pattern. For example, in species of the pulchra group which have been studied in detail, the males possess a ventrosublateral pair of cervical setae just posterior to the amphid; character d, Fig. 4. In the armata group, there are bilaterally symmetrical pairs of elongated setae, character a, Fig. 4 (e.g. S. elongata: Platt, 1984). It may be that the shorter cervical setae in species of the other groups may also prove to have a specific pattern, but they have not yet been studied in sufficient detail. The cervical setation patterns are considered autapomorphic features of the pulchra and armata groups. Amphids: The amphids in all Sabatieria species are spiralled ventrally to the centre. As with the Ethmolaimidae (Platt, 1982: 190) the exact turns counted can vary within species. The turns always begin on the dorsal side but it is the last inner part of the spiral that can vary. However, in this work, the amphids are simply denoted by the number of complete dorsal quadrants, as shown in Fig. 7. In this way, an amphid with 2$, 2$, 2$, or 3 turns are all referred to as 3 turns. Ninety per cent of Sabatieria species have 3 turns (Fig. 5A), some have 4 turns (Fig. 5D), but only one species, S. bonessa, unequivocally has 2 turns (Fig. 13). The number of turns is not congruent with any other Sabatieria character states and the widespread occurrence of similar amphids indicates the feature is plesiomorphic. Copulatory structures: All Sabatieria species have paired spicules of equal size and a gubernaculum, features which are of course plesiomorphic. All valid species also have paired apophyses attached to the gubernaculum, a feature which is found in all but four comesomatid genera and in several other nematode taxa, particularly within the Monhysterida. However, as discussed earlier, the hypothesis supported here is that comesomatids are part of the Chromadorida: the consequence being that, unless apophyses are plesiomorphic for all nematodes, then they must have evolved on more than one occasion and the character is not congruent. Those comesomatid genera lacking apophyses must have lost them secondarily according to the principle of commonality. The apophyses in other genera of the Comesomatidae and in most Sabatieria species are dorso-caudally directed and straight (Fig. 8E, G). However, in some Sabatieria species, the apophyses are characteristically curved (Fig. 8A), a synapomorphic feature for celtica and ornata groups; character f, Fig. 4. The median piece (cuneus) of the gubernaculum in the Pulchra group typically has a well-developed caudally directed bar extending from its dorsal end (Fig. 8B, E), an apparently unique feature interpreted as apomorphic; character c, Fig. 4. The detailed structure of the spicules of Sabatieria species reveals the presence in several species of a central cuticularized internal projection from the proximal end (Fig 8B, D, G) of varying length which may (Fig. 12, S. stekhoveni) or may not (Fig. 8A-D) be continued as a central lamella or list. Proximal projections and central lists may be found in several outgroups (e.g. Spirinia parasitifera BMNH: , Pomponema sedicima BMNH: ) and so must be considered plesiomorphic. Sabatieria species with elongated narrow spicules

14 H. M. PLATT A 2 1 I1 / If s 3 2 I!..!. I I l l 2; I Figure 7. Explanation of amphid-type assessment. A, 2 turn amphids; B, 3 turn amphids. C, 4 turn amphids. s in row A indicates where counting of the number of turns begins and the cross divides the spiral into quarters. The number beneath each drawing indicates the number of turns exact to the nearest quarter. See text for further explanation. often lack any central structures (Fig. 8F), as occurs in other comesomatids (e.g. Dorylaimopsis punctata BMNH: ). Sabatieria species typically possess a ventral series of precloacal supplements which in most species are represented by indistinct pores which are often difficult to observe (Fig. 21H; S. praedatrix). It is probable that they have simply been overlooked in some species reported devoid of supplement. The possession of rows of supplements in comesomatids and other chromadorids suggests their presence is a plesiomorphic feature. Lorenzen (1981) pointed out that many comesomatids have slightly arched supplements of a kind which do not appear elsewhere in the Chromadoria. With the possible exception of Scholpaniella pontica (Fig. l), which in any case is a problematical poorly described species, this precise form of conspicuous supplement appears to be confined to certain groups of Sabatieria species alone, namely the pulchra (Fig. 5F), celtica (Fig. 5C) and ornata (Fig. 5H) groups; character b, Fig. 4. Consequently, this has been interpreted as a synapomorphic feature within Sabatieria establishing the holophyly of the three combined species groups. The precise distribution of these well-developed supplements appears to fall into three patterns, two of which had previously been identified by Jensen (1981):

15 REVISION OF SAHA TIERI.4 Figure 8. Copulatory apparatus. A, S. ornata; B, S. brevlseta; G, S. praedatrix. Scale bar = 10 prn. C-E, S. pulchra; F, S.!yonessa;

16 12 H.,M. PLATT (1) equally spaced or gradually increasing in distance apart anteriorly, the celtica group (Fig. 5C); (2) a posterior group of more closely situated supplements, separated from the rest (character g, Fig. 4), the ornata group (Fig. 5H); (3) an anterior group of more closely situated supplements (character e, Fig. 4), the pulchra group (Fig. 5F). Compared with the distribution of supplements in the praedatrix and armata groups, the celtica condition would appear to be plesiomorphic, with the ornata condition a derivation from it. This would imply that the pulchra group is the apomorphic sister-group of celtica + ornata. The unique supplement disposition in both the ornata and pulchra groups are autapomorphic features establishing the holophyly of each. All Sabatieria species appear to have a ventral precloacal spine (Figs 20E, 21F), although it appears to have been overlooked according to some of the figures in the literature. This is a plesiomorphic character known to occur in several of the outgroups studied (e.g. Ethmolaimidae, Chromadoridae, Cyatholaimidae). Reproductive system: All Sabatieria species studied had two opposed outstretched gonads. The phylogenetic interpretation of the outstretched condition of the female ovary has been discussed above. In most species studied the anterior ovary was to the left, the posterior to the right of the gut: in S. kelletti and S. punctata it was vice versa. In all species studied, the anterior testis was to the left, the posterior to the right of the gut (Fig. 5G). This consistent leftlright disposition of gonads relative to the gut is plesiomorphic for Sabatieria according to Lorenzen (1981). The condition in S. kelletti and S. punctata, in which the ovaries are right/left and the testes left/right, is unusual, according to Lorenzen ( ). TAXONOMIC REVISION AND IDENTIFICATION In the following diagnoses, only those important features which serve to distinguish the taxon in question from all others of equivalent rank are given: homologous characters shared by other taxa are usually omitted. In this way, an attempt has been made to avoid an extended list of mixed primitive and derived characters which provide more of a morphological description than a taxonomic diagnosis. *rwe GENUS: Comesoma Bastian, Family Comesomatidae Filipjev, DIAGNOSIS: Punctated cuticle. Spiral amphids. Precloacal supplements. Outstretched ovaries. Paired apophyses on gubernaculum. TYPE GENUS: Sabatieria Rouville, Subfamily Sabatieriinae Filipjev, 1934 DIAGNOSIS: Comesomatidae. Posterior part of buccal cavity not expanded. Spicules not elongated.

17 REVISION OF.SABATIERfA 4:3 REMARKS: No derived characters uniquely shared by genera of this subfamily could be found: the characters of the diagnosis are not synapomorphies and the taxon cannot be considered holophyletic. Setosabatieria gen. nov. TYPE SPECIES: Sabatieria hilarula De Man, DIAGNOSIS: Sabatieriinae. Cuticle not punctated. Sublateral rows of long cervical setae. REMARKS: Only the presence of long cervical setae is autapomorphic, the absence of cuticle punctations being shared with Laimella and Cervonema. However, the characters of the diagnosis serve to separate the new genus from Sabatieria. The erection of the new genus is based on a study of S. hilarula (see below), but Sabalieria jibulata Wieser, 1954 also appears to conform to the diagnosis of Setosabatieria and is therefore transferred accordingly. Selosabatieria consists of the following valid species: S. Jibulata (Wieser, 1954) syn. nov.; S. hilarula (De Man, 1922) syn. nov. The species may be distinguished using the information in Table 1 and Fig. 9. Note: In the following species revisions, a synopsis is given of descriptions in the literature in addition to new observations, where appropriate. Where known, the number of specimens actually used in each description is given, not the total number of specimens reported to be found in an ecological survey. B Figure 9. Species of Setosabatieria. A, S. hilarula; B, S. fibulala. Scale bars: heads = 10 prn; tails = 30 prn.

18 ~~ 44 H. M. PLATT Table 1. Differentiating data of male Setosabatieria species Species TL a Hd A% At R3 Cs Spic Ps T S. fibulutu ~ S. hilarulu Abbreviations: TL, total body length (pm) to nearest 5 pm; a, De Man ratio a (total body length i maximum body diameter); Hd, head diameter as percentage of posterior oesophagus body diameter; AO/,, amphid diameter as percentage of corresponding body diameter; At, number of turns of amphid; R3, R3 sensilla length as percentage of head diameter; Cs, number of cervical setae in each sublateral row; Spic, spicule chord length as proportion of cloacal body diameter; Ps, number of precloacal supplements; T, tail length measured in cloacal body diameters. Note: Unless stated otherwise, in this and subsequent tables, the data have been extracted from all available descriptions, including additional information from specimens studied here. Setosabatieria hilarula (De Man, 1922) (Figs 9, 10, 11A-C; Table 1) Sabatieria hilarula De Man, Sabatieria chitwoodi Wieser, Syn. nov. Sabatieri Sabatieria scotlandia Inglis, Syn. nov. jubata sensu Weiser, 1954 nec Cobb, Syn. nov. Material. 6 dd, 5 99, 5 juveniles from kelp holdfasts off west coast Scotland (types of S. scotlandia) BMNH , 1 $! from Tamar estuary, SW England (leg. R. M. Warwick) BMNH Jd from Northumberland coast, NW England (leg. R. M. Warwick) BMNH , 1 $! from Bresund, Denmark (specimens previously described by Jensen, 1979a). Descriptions. Several good descriptions available (see Gerlach & Riemann,. 1973), the most recent being Jensen (1979a). Additional observations: Very faint external striations can be discerned which become more widely spaced posterior to the oesophageal region and are also discontinuous laterally (Fig. 9). The Scottish specimens do not appear to be punctated, in contradiction with Inglis ( 1961). The gubernaculum region is characteristically obscured by lateral glandular structures (rectal/cloacal glands?). Supplements are minute and difficult to detect so counts are subject to possible errors. However, in the Scottish specimens, three had 13, 14 and 15 supplements each (Inglis, 1961 stated that there were about 12), in the Tamar specimens there were 15 supplements and in the Northumberland specimens 14, 15 and 16. In the Scottish specimens, there are two gubernaculum apophyses, not one as implied by Inglis (1961). Lateral to the cloaca are usually observed leaf-like extensions of the cuticle (Figs IOC, 11A-C): function unknown. In all males examined, bundles of thread-like structures were observed lying subventrally either side of the vas deferens (Fig. 10D): the threads or fibres appeared to run posteriorly from the body wall to the vas deferens. Gonads: anterior left, posterior right of gut. Remarks. Sabatieria scotlandia Inglis, 1961 was not included in Jensen s (1979a) revision. However, re-examination of the type material showed the specimens to be identical with the specimens described by Jensen (1979a): the two species are therefore synonymized. Jensen s specimens had cervical setae per row:

19 REVISION OF SABATIERIA 4 5 Figure 10. Sefosubufieria hilurulu. A-B, head of d from Bresund, showing amphid (A) and R1, R2 and R3 cephilic sensilla, arrowed (B); C, cloaca1 region of 3 from Tamar estuary, showing leaf-like cuticle extensions (arrow); D, thread-like structures lying lateral to vas deferens, J from Tamar estuary. Scale bar = 10 pm. Inglis had These observations extend the known range in cervical setae number in this species to 3-16 per row. Jensen (1979a) reported that in his 81, the right spicule was more curved than the left. This appears to be due to the specific orientation of the specimen and a re-examination of the material indicated that the spicules were more or less identical. As regards the thread-like structures observed attached to the vas deferens, I am unaware of them having been reported before. Jensen (1979a) discussed the similarity between S. hilarula, S. chitwoodi and S.jubata. According to Jensen, S. chitwoodi only differs from S. hilarula in the length of cephalic and cervical setae, and in having a shorter cylindrical portion

20 46 H. M. PLATT Figure 1 I. A-C, Sefarabafieria hilarula: copulatory apparatus of three 88 from Scotland; D, posterior region of Sabaticria praedatrix. Scales bar: A-C = 10 pm; D = 30 pm. of the tail. In other extant descriptions of S. hilarula (see Gerlach & Riemann, 1973) these characters show considerable variation so it is proposed that the two species be synonymized. Likewise, S. jubuta sensu Wieser 1954 only differs from S. hilarula in the number of cervical setae, but since the range in S. hilarula (including S. scotlandia) is 3-16 per row, then to keep S.jubata separate for its rows of 21 seems unreasonable. Jensen (1979a) pointed out that the relative measurements of the amphids were different, but Wieser (1954) was uncertain of the size of the male amphid and the figure given is of the female. Therefore, S.jubata sensu Wieser, 1954 can be considered synonymous with S. hilarula. S. jubata Cobb, 1898 is inadequately described and here considered dubious. Records. 33: worldwide. Setosabatieria Jbulata (Wieser, 1954) (Fig. 9; Table 1) Description. Wieser (1954), type. 1 d, Cuticle ornamentation indistinct. 3-4 cervical setae in 6, 6-7 in 99. Male amphid 4.25 (= 5) turns; female 3.75 (= 4) turns. Tail with short cylindrical part. Sublittoral fine sand, m depth, Chilean coast.

21 REVISION OF SARA71EKIA 47 Remarks. Setosabatieria jbulata is characterized most easily by the number of 3 amphid turns and tail shape. Records. 1: Chile. Sabatieria Rouville, 1903 TYPE SPECIES: Sabatieria celtensis Rouville, DIAGNOSIS: Sabatieriinae. Cuticle punctated, without longitudinal files of dots. R3 sensilla longer than R2 sensilla and not situated close together. Amphid situated immediately posterior to R3 sensilla. REMARKS: No derived characters uniquely shared by species of this genus could be found: the characters of the diagnosis are therefore a rather unsatisfactory combination of plesiomorphic characters which simply serves to distinguish Sabatieria from other genera of the subfamily. This lack of derived characters, both for the genus and the subfamily, suggests that Sabatieria may represent the primitive form from which all other comesomatids may be derived, especially if abundance of species and frequency of occurrence is taken to be an indication of the primitive state, as discussed earlier. One species, S. microseta Timm, 1967, has very small R3 sensilla, small buccal cavity, irregular lateral cuticle punctations and no precloacal supplements. These features conform to the generic definition of Pierrickia Vitiello, 1970 and the species is transferred accordingly. Of the remaining species, only 36 are considered valid and the identification of the species assigned to each of the five subgroups will first be discussed. This is followed by an alphabetical review of each valid species and then brief remarks on the eight species considered inquirendae. Sabatieria subgroups (Fig. 4) Praedatrix group: The 17 species included in this group (Table 2) are those with simple tubular or pore-like supplements (usually about 17 in number) and straight gubernaculum apophyses. As the group is currently constituted, none have spicules with central lists separated from the proximal projection and most have lateral cuticle differentiation of larger, more widely spaced dots. Most have 3 turn amphids except S. lyonessa (2 turns) and S. triplex (4 turns). However, no au tapomorphies could be found. The species may be distinguished by careful consideration of the information in Table 2 and Figs 12 and 13. Most species are poorly known and have been infrequently reported: 8 species recorded only once, 4 twice, 3 three times and 1 five times (S. granger). The most frequently reported species is S. praedatrix ( 14 times), although five of these were by A1lgt.n and should be treated with some degree of caution. Armata group: The 5 species (Table 3), like those of the praedatrix group, have simple tubular supplements; they were probably erroneously not reported present in S. supplicans. They differ from the praedatrix group and other species in having characteristically elongated R3 cephalic ( h.d.) and cervical sensilla, and slender bodies ( a ) usually > 65, except S. migrans).

22 48 H. M. PLATT 5. praedafrix S. intermisso S. granifer \ S. sfekhove. I S. laws1, S. poradoxa L I S. vasico/a., S triplex S. paracupida Figure 12. Sabatieria species belonging to the praedatrix-group (continued in Fig. 13). Sabatieria intermissa, S. granger, S. triplex after Wieser (1954); S. stekhovcni, S. vasicola after Vitiello (1970); S. paradoxa, S. paracupida after Wieser & Hopper (1967); S. pruedatrix, S. laws; original. Numbers = supplements. Scale bars as in Fig. 9. Sabatieria strigosa and S. macramphis outside the armata group have R3 sendla as long but the long cervical sensilla appear to be unique (autapomorphic). All have 3 turn amphids except S. longispinosa (4 turns). The species may be distinguished using the information in Table 3 in conjunction with the pictorial key (Fig. 14). Sabatieria armata and S. longispinosa have been recorded 4 and 5 times respectively, the rest only once. Pulchra group: The 6 species (Table 4) are distinguished by the disposition of the precloacal supplements, which are usually conspicuous and relatively few in

23 REVISION OF SARATIERIA S. fyonessa S. conicouda s. 0Iafa \ \ S. fatcifera S. parabyssatis S. oncudlono Figure 13. Further Sabatierza species belonging to the paedatrix-group. Sabatieria lyonessa after Warwick (1977); S. conicauda after Vitiello (1970); S. alata after Warwick (1973); S. demani after Stekhoven & Bresslau (1940); S. dodecaspapillata after Kreis (1929); S. fakifera, S. parabyssalis, S. ancudiana after Wieser (1954). Numbers = supplements. Scale bars as in Fig. 9. number (5-9), by the characteristic gubernaculum median pieces and short paired cervical setae: all appear to be autapomorphies. They also tend to be shorter and stouter than other Sabatieria species. Sabatieria breuiseta and S. punctata have 4 turn amphids, the rest have 3 turns. The species may be distinguished using the information in Table 4 in conjunction with the pictorial key (Fig. 15). Sabalieria pulchra and S. punctata are by far the most commonly reported species (about 40 and 2 1 times respectively). Sabatieria breviseta has been reported 5 times, S. mortenseni and S. pissina both twice and S. propissina once. Celtica group: The 4 species (Table 5) all have curved gubernacular apophyses, 4

24 50 H. M. PLATT Table 2. Differentiating data of male Sabatieria praedatrix group species Species TL a Hd A% R3 Spic Ps T S. alata S. ancudiana S. conicauda S. demani S. dodecaspapillata S. falcqera S. granif r S. intermissa S. lawsi S. gonessa S. parabyssah S. paracupida S. paradaxa S. praedatrix S. stekhoveni S. lriplex S. uasicola -~ ~ Abbreviations: see Table I c c c ~13-17 ~ ? conspicuous supplements (?S. furcillata), 3 turn amphids, and tend to be relatively large and stout but there are no apparent autapomorphies. They may be distinguished using the information in Table 5 in conjunction with the pictorial key (Fig. 16). With about 40 records, S. celtica is by far the commonest species: S. furcillata has been found twice and the other two once only. Ornata group: The 4 species (Table 6) are generally similar to species of the celtica group except for the presence of a posterior group of more closely situated precloacal supplements, considered an autapomorphic feature. All have 3 turn amphids, except S. ornata, which has 4 turns, although it may easily be mistaken for 3 turns. The species may be distinguished using the information in Table 6 in conjunction with the pictorial key (Fig. 17). Sabatieria ornata is the most frequently reported (14 times) followed by S. abyssalis (8 times): both are very similar. Sabatieria longiselosa and S. macramphis have been found twice and once respectively. Valid species In each of the following reviews, where possible the number of specimens on which the descriptions were based is given-which may be different from the total number of specimens recorded from a locality although the distinction is not always clear. Unless specifically recorded that material was available, infarmation is derived from the published descriptions. Records and localities are as in Gerlach & Riemann (1973): only later records are referenced here. See Gerlach & Riemann (1973) also for full synonymy. Sabatieria abyssalis (Filipjev, 1918) (Fig. 17; Table 6) Descriptions. (1) Filipjev (1918), type. 1 6, 1 0: total length (TL) = 1650, 1500 pm; a = 33, 25. Cuticle punctations irregular, not forming annules.

25 REVISION OF SABATIERIA 5 I S. ormoto 5. long/splnoso S. elongoto 5. mfgrons {icons Figure 14. Sabatieria species belonging to the armata-group. Sabatieria armata after Gerlach (1952); S. migrans after Jensen & Gerlach (1977); S. supplicans after Gerlach (1956); S. longispinosa original except for amphid after Riemann (1966); S. elongata original. Numbers = supplements. Scale bars as in Fig. 9. Table 3 Differentiating data of Sabatieria armata group species Species TL a Hd A% R3 Spic Ps T S. armata S. alongata S. longispinosa * S. migrans S. supplicans ? 8 Abbreviations: see Table I *Corpus gelaturn usually uncoiled: figure taken from Riemann (1966)

26 ~~ 52 H. M. PLATT S. pulchra S. morfenseni S. punctoto S. breviseta S. propissino S mssino fid 1... ':.:;:;.... ** :>:*...., c :,[\I/ Figure 15. Sabaticria species belonging to the pulchm-group. Sabaficriu martensmi after Ditlevsen (1921); S. propissina after Vitiello (1976); S. pzssinn after Vitiello (1970); others original. Numbers = supplements. Scale ban as in Fig. 9. Table 4. Differentiating data of male Subutieriu pulchru group species Species TL a Hd A% R3 Spic Ps T S. breviscta S. mortensmi c o S. pissina ? S. propitsinn ? S. pulchra S. pundata Abbreviations: see Table I

27 REVISION OF SABATIERIA S. celtica 5'. kelletti \ \ S. furcillota \ I Figure 16. Sabatieria species belonging to the celtica-group. Sabatieria strigosa after Lorenzen ( 1972); S. furcillata after Wieser (1954); others original. Numbers = supplements. Scale bars as in Fig. 9. Table 5. Differentiating data of male Sabatieria celtica group species Species TL a Hd A% R3 Spic Ps T S. celtica S. furcillata S. kelletti S. stripsa ? Abbreviations: see Table 1 Amphid 2+ turns in figure, 3+ in text. Male tail 4.5 a.b.d. in figure, 5.3 in text. Four posterior supplements closer together. Mussels and mud, Black Sea. (2) Timm (1961). 1 8: TL = 1880 pm; a = Tail with a slightly longer cylindrical part (55% of total). No mention of disposition of supplements.

28 54 H. M. PLATT 5. ornafa 5. abyssalis S. macramphis S. long/sefosa Figure 17. Sabatieria species belonging to the omata-group. Sabatieria ornata after Jensen (1979a); S. abyssalis after Filipjev (1918); S. macramphis after Lorenzen (1972); S. longbetosa after Kreis (1929). Numbers = supplements. Scale bars as in Fig. 9. Table 6. Differentiating data of male Sabatieria ornata group species Species TL a Hd A% R3 Spic Ps T S. abyssalis S. longisetosa S. macramphis ? S. omata Abbreviations: see Table 1 Otherwise, similar to Filipjev (1918). Intertidal mud, Sonadian Island, Bay of Bengal. (3) Groza-Rojancovski (1973). 1 3: TL = 1850 pm; a = 28. Similar to Filipjev ( 1918) except gubernaculum apophysis, which is depicted as unusually short for Sabatieria species m depth mud, Rumanian coast.

29 REVISION OF SABATIERIA 55 Remarks. This species is remarkably similar to S. ornata (Ditlevsen, 1918) as redescribed by Jensen (1979a). From the available descriptions in the literature, the only significant difference appears to be that S. ornata has a 4 turn amphid whilst S. abyssalis has 3. Records. 8: Black Sea (4: Groza-Rojanovski, 1973; Sergeeva, 1976); Mediterranean (1); Bay of Bengal (3). Sabatieria alata Warwick, 1973 (Figs 13, 18A-D; Table 2) Material. 1 3, 1 0: BMNH Descriptions. (1) Warwick (1973), type. 2 d&, 2 99: TL = pm; a = About 21 inconspicuous supplements. Mud at 3147 m depth, off Muscat and Oman, Arabian Sea. (2) Additional observations. Lateral cuticle punctations in transverse rows in 9 and midbody of 3 (Fig. 18C) but more irregular in postamphid and caudal regions of 6 (Fig. 18B, D). Amphid 3 turns (Fig. 18A). Conspicuous cloaca1 glands present. Anterior gonad left, posterior right of gut. Remarks. The most distinguishing feature of Sabatieria alata appears to be the characteristic bend at the distal end of the spicule (Fig. 13). Records. 1: Indian Ocean. Sabateiria ancudiana Wieser, 1954 (Fig. 13; Table 2) Description. Wieser (1954), type. 7 dd, 8 99: TL = pm; a = (mean = 58). Fine sand, 39 m depth, Chilean coast. Remarks. Sabatieria ancudiana has no unique distinguishing features. Records. 3: Chile (1); North East coast U.S.A. (2: Tietjen, 1977, 1980: Tietjen s record of Sabatieria anucaudiana from Long Island sand is assumed to be S. ancudiana). Sabatieria armata Gerlach, 1952 (Fig. 14; Table 3) Description. Gerlach (1952, type and 1953). 2 63, 3 99: TL = pm; a = According to Gerlach s (1952) description of a female, the cuticle has irregular punctations, larger laterally than medially although the figure shows the punctations in transverse rows. From Gerlach s (1953) figure of the male, the 9 supplements extend some 120 pm from the cloaca, which is about 4.9% of the total length (TL) or 5.2% of the TL minus tail length (TL ). Intertidal sand, Italian coast. Remarks. Sabatieria armata is distinguished from others of the group by the distinctive distal spicule tip (Fig. 14). There is some confusion about the form of the cuticle ornamentation but, for the purposes of this analysis, I will assume it

30 Figure 18. A-D, Sabatieria alata. A, Amphid; B, cuticle in posterior oesophagus region; C, cuticle in midbody region; D, cuticle in caudal region; E-G, S. celtica from Strangford Lough, Northern Ireland; E, cuticle pattern posterior to amphid; F, cuticle in midbody region; G, cuticle in cloaca1 region. Scale bar = 10 pm.

31 REVISION OF SABATIERIA 57 is similar to S. elongala; namely, with larger lateral dots more or less arranged in transverse rows. Records. 4: Mediterranean (2); New England, U.S.A. (1); NE Ireland (1). Sabatieru breviseta Stekhoven, 1935a (Figs 5E, F, J, 9B, 15; Table 4) Sabatieria quadripapillata sensu De Coninck & Stekhoven, 1933 nec Filipjev, Material. 3 63, 2 99: BMNH Descriptions. (1) De Coninck & Stekhoven (1933), type. 19 only: TL = 1480 pm; a = 34. Amphid 2+ turns (55% c.d.). Intertidal sand and shells, Belgian coast. (2) Stekhoven (1935a). 1 d, TL 8 = 1140 pm; a = 36. Male amphid 4 turns (75% c.d.). Cuticle with fine uniform punctation without differentiation of the lateral field. Fine sand, Belgian coast. (3) Platt (1984). 3 S6, 2 99: TL = pm; a = Amphids 3; (almost 4) turns in Sd (80-85y0 c.d.) and 2+ (almost 3) turns in 99 (60% c.d.). Gonads: anterior left, posterior right of gut. Bristol Channel. Remarks. Stekhoven (1935a) only reported 5 supplements but he may have overlooked the most posterior one: in his figure, the first supplement is situated well anterior to the proximal end of the spicule. Sabatieria breviseta belongs to the putchra group and is distinguished by cuticle pattern, large amphids of 4 turns (8) and prominent gubernaculum median piece. Records. 5: North Sea (4); Bristol Channel (1). Sabatieria celtica Southern, (Figs 6E, 16, 18E-G; Table 5) Sabalieria cupida Bresslau & Stekhoven in Stekhoven, 1935b Sabalieria longiseta S teiner, Sabatieria tenuicaudata sensu Stekhoven, 1943 nec Bastian (1865) Purasabatieriu longiseta Allgkn, 1934 Material. Several 68 and 99: BMNH ; ; ; Descriptions. Several descriptions are available of this well-known species (see Gerlach & Reimann, 1973) among the most recent being Lorenzen (1972) and Platt (1984). Type locality: sand and shells at 40 m depth, Clew Bay, W Ireland. Remarks. This very commonly reported species appears to display rather a wide range in the intraspecific variation of certain character states, especially the relative R3 sensilla length (Table 5). Records. 40: worldwide distribution, although those from non-european waters should perhaps be treated with some caution.

32 58 H. M. PLATT Figure 19. Subatieria lyonessu. A, Anterior showing amphid and lateral cuticle ornamentation; B, cuticle in midbody region; C, cuticle in cloaca1 region; D, entire 0; E, $. posterior region; F, precloacal supplements. Scale bars: A-C, F = 10 pm; D = 300 pm; E = 100 pm.

33 REVISION OF SABATIERIA 59 Sabatieria conicauda Vitiello, 1970 (Fig. 13; Table 2) Description. Vitiello (1970), type. 2 33, 1 0. Sublittoral mud, French Mediterranean coast. Remarks. Vitiello (1970) considered the species to be characterized by the caudal punctation: Les puntuations sont plus grosses et consistant en tubes assez ilevts et protminents. However, how this is very different is not clear to me from Vitiello s figure. Jensen (1979a) considered that the description was based on specimens with broken tails, but since Vitiello (1970) had 13 specimens available it seems to me unlikely that they all had broken tails. Apart from the tail, S. conicauda would appear to belong to the praedatrix group. Records. 2: Mediterranean (Vitiello, 1976). Sabatieria demani Bresslau & Stekhoven in Stekhoven, 1935b (Fig. 13; Table 2) Description. Stekhoven (1935b), type. Bresslau & Stekhoven (1940), description of same specimens. 1 3, 1 9: TL = 8 mm (both 3 and 9); a = 63,60; c = 25,29 Male amphid 3 turns, 65% c.d.; female amphid 2 turns, 37% c.d. Pobgoridiusground, near Helgoland, North Sea. Remarks. Sabatieria demani would appear to be like a very large S. celtica, as pointed out by Bresslau and Stekhoven (1940) themselves. The wider definition of S. celtica (see above) would now include two of Bresslau and Stekhoven s points of separation: amphid and R3 sensilla size. Apart from total body length, the features still separating S. demani from S. celtica (and others) would appear to be the sexual dimorphism of the amphid, detailed structure of the spicules and gubernaculum and possibly relative length of the tail (c-ratio compared with normal range of about 10-18). Incidentally, the figures in Stekhoven (193513) were wrongly labelled; fig. 232A is the $2 head (not 3) and fig. 232B is 8 tail (not 9). Records. 1: North Sea. Sabatieria dodecaspapillata (Kreis, 1929) (Fig. 13; Table 2) Description. Kreis (1929), type. 33: TL = pm. English Channel. Remarks. Kreis (1929) could not distinguish punctations in this species: doch nicht in Punkte auf geloste Ringelung. I interpret this as being very fine regular punctations such as those of S. breviseta. Apart from this, S. dodecaspapillata is only separated by very minor differences from others of the praedatrix group, such as S. abyssalis. North coast Brittany, France. Records. 1 : English Channel.

34 60 H. M. PLATT Material. 2 dc3: BMNH Sabatieria elongata Jayasree & Warwick, 1977 (Figs 6B, 14; Table 3) Descriptions. (1) Jayasree & Warwick (1977), type. 2 d& TL = 3430, 3575 pm. Intertidal sand, Firth of Clyde, Scotland. (2) Platt (1984), re-examination of types. Testes: anterior left, posterior right of gut. Remarks. Sabatieria elongata belongs to the armata group but is distinguished by combinations of spicule structure, supplement number, tail shape and cuticle ornamentation. Records. 1: W Scotland. Sabatieria falcqera Wieser, 1954 (Fig. 13; Table 2) Description. Wieser (1954), type , Cuticle with transverse rows of dots, larger laterally, sometimes irregularly arranged. Male amyt3hid 2$-3 turns, female amphid 2+-2$ turns. About 10 supplements. Sublittoral fine to coarse sand, m depth, Chilean coast. Remarks. This is one of several identikit species, practically indistinguishable morphometrically. However, S. falcifera can be distinguished by careful consideration of the cuticle ornamentation, copulatory apparatus and tail shape. Records. 2: Chile (1); New England, U.S.A. (1: Tietjen, 1969). Sabatieria furcillata Wieser, 1954 (Fig. 16; Table 5) Description. Wieser (1954), type. 44 dd, Cuticle: 3 transverse rows of dots per annule; lateral dots slightly larger and horizontally further apart. Sublittoral clay, mud and coarse sand, m, Chilean coast. Remarks. Relatively distinct cuticle annulation would appear to distinguish this member of the celtica group but it must be borne in mind that clarity of annulation can vary among specimens. Records. 2: Chile (1); North Carolina coast, U.S.A. (1: Tietjen, 1976). Sabatieria granger Wieser, 1954 (Fig. 12; Table 2) Sabatieria granulosa Vitiello & Boucher, Syn. nov. Description. Wieser (1954), type. 7 33, Lateral differentiation of cuticle consists of larger irregularly arranged dots, especially noticeable in cervical and caudal regions. Sublittoral clay to coarse sand, m depth, Chilean coast. Remarks. The species is characterized by the coarse lateral punctuations. The figure of the male tail provided by Wieser (1954) omits the distal part, so in the pictorial key I have completed it with part of Wieser s illustration of the female

35 REVISION OF SABA TIERIA 61 tail. Vitiello & Boucher s (1971) species, S. granulosa, appears to be morphometrically identical to S. granger: Vitiello & Boucher suggest there is a difference in the detailed structure of the copulatory apparatus, but when allowance is made for possible different orientation and interpretation, they appear to be quite similar. On this basis, I propose to synonymize the two species. Records. 5: Chile (1); Mediterranean (4: Boucher, 1972, 1973; Vitiello, 1974, 1976b). Sabatieria intermissa Wieser, 1954 (Fig. 12; Table 2) Description. Wieser (1954), type. 8 Jd, Cuticle: fewer lateral rows of dots which are rod-shaped in mid-body. Intertidal and subtidal sand, Chilean coast. Remarks. The species is remarkably similar to S. praedatrix De Man, 1907a except for the larger spicules, differently shaped gubernacula and larger male amphid. Records. 1: Chile. Sabatieria kelletti Platt, 1983 (Figs 5A, B, C, 6F, 16; Table 5) Material. 6 88, 1 9, 5 juveniles: BMNH Description. Platt ( 1983). Remarks. Sabatieria kelletti belongs to the celtica group. Its distinguishing characteristics are the shape of the head, presence in the male of a ventral caudal swelling and possibly the three characteristic subventral cervical setae. Records. 1: Antarctic peninsula. Sabatieria lawsi Platt, 1983 (Fig. 12; Table 2) Material. 6 83, 6 99, 19 juveniles: BMNH Description. Platt ( 1983). Remarks. Sabatieria lawsi belongs to the praedatrix group with widely-spaced simple precloacal supplements and fairly straight gubernaculum. It is very similar S. zotieseri nom. nov. (= S. heterura sensu Wieser, 1954 nec Cobb, 1898) but may be distinguished by the detailed structure of the copulatory apparatus. Records. 1 : Antarctic peninsula. Sabatieria longisetosa (Kreis, 1929) (Fig. 17; Table 6) Description. Kreis (1929), type. 3 d8. Cuticle punctations not recorded supplements, posterior 4 closer together. North Coast of Brittany, France. Remarks. This species belongs to the ornata group, being morphometrically similar to S. ornata itself except for having longer cephalic setae: the tail is also a

36 62 H. M. PLATT different shape and the amphid appears to have half a turn less. However, Kreis' (1929) figure of the head is rather sketchy and a redescription is needed. Records. 2: English Channel (I), Scilly Isles (1: Warwick & Coles, 1977). Sabatieria longispinosa Lorenzen, 1972 (Fig. 14; Table 3) Material. 2 dd and 2 $29 from NE Ireland: BMNH dd, 1 $? from Bresund, Denmark (type locality): BMNH Descriptions. (1) AllgCn ( 1935), type. Poor description based on female only. (2) Riemann (1966). 2 dd, (3) Platt (1984). 5 d~3, Gonads: anterior left, posterior right of gut. Remarks. Sabatieria longispinosa belongs to the armata group and is distinguished by the larger male amphid, which normally has the corpus gelatum uncoiled, the shape of the male tail and possibly by the cuticle ornamentation. Records. 5: North Sea (2); Bresund (2: Platt, 1984); Northern Ireland (1 : Platt, 1984). Sabatieria lyonessa Warwick, 1977 (Figs 8F, 13, 19; Table 2) Material. 1 d, 1 0, BMNH Descriptions. (1) Warwick (1977), type. 1 6, 1 0. Laminaria holdfasts, Scilly Isles. (2) Additional observations. Lateral cuticle has fewer transverse rows of dots (Fig. 19B) which may appear somewhat longitudinally elongated in midbody. Gonads: anterior left, posterior right of gut. Remarks. Sabatieria lyonessa appears to be most closely associated with the praedatrix group of species. It is distinguished by the small size of the amphid (Fig. 19A), tail shape (Fig. 19E) and number of supplements. Records. 1: Scilly Isles. Sabatieria macramphis Lorenzen, 1972 (Fig. 17; Table 6) Description. Lorenzen (1972), type Sublittoral fine sand, 25 m depth, near Helgoland. Remarks. This species is rather like S. ornata, except the R3 sensilla are larger and there are more supplements. Records. 1: North Sea. Sabatieria migrans Jensen & Gerlach, 1977 (Fig. 14; Table 3) Descripion. Jensen & Gerlach (1977), type , Transverse rows of dots which in the lateral field are coarser and, at least in the cloaca1 region,

37 REVISION OF SABATIERIA 6 :3 there are fewer rows. Cloaca1 (rectal) glands present indistinct supplements; cannot be counted with certainty. Intertidal medium sand, Bermuda. Remarks. Sabatieria migrans belongs to the armata group. According to Jensen & Gerlach s (1977) two figures of the male head, there appear to be characteristic lateral and sublateral cervical setae situated just posterior to the amphid. The species is distinguished from others of the armata group by the larger number of supplements. Records. 1 : Bermuda. Sabatieria mortenseni (Ditlevsen, 192 1) (Fig. 15; Table 4) Descriptions. (1) Ditlevsen (1921), type. 1 8, 1 9. Six supplements, the anterior four being closer together. Could not detect any cuticle patterns even with immersion lens. (2) Pastor de Ward (1984): see note added in proof. Remarks. This species is similar to S. breviseta but it has much longer R3 sensilla. Allgtn (1947) claimed to have found a single female of the species in California, but the description is poor and the record should be considered dubious. Allgh s (1959) report of the species from S Atlantic waters should be treated with caution. Records. 2: Auckland Islands (1); Brazil (1). Sabatieria ornata (Ditlevsen, 19 18) (Figs 5G-I, 8A, 17, 20; Table 6) Sabalieria proatpssalis Vitiello & Boucher, Syn. nov. Sabatieria similis (Allgen, 1933). Syn. nov. Material from NE coast England, BMNH Descriptions. (1) Ditlevsen (1918), type. 1 $. TL = 2200 pm; a = 47. No R3 sensilla observed. 14 supplements; 5 posterior ones closer together and set-off from the others. Sublittoral shell-ground, about 30 m, Bresund, Denmark. (2) Stekhoven (1946) R3 sensilla detected, 52% h.d. Sublittoral m depth, Skagerrak, Sweden. (3) Jensen (1979a) , from 0resund, Denmark (sublittoral sand, m depth) and re-examination of type material. (4) Additional observations. Lateral cuticle pattern generally appears irregular but may occasionally resemble anastomosing rows (Fig. 20C). Of material studied, six had 11 supplements and the others 10, 12 and 13 each. Eight had the 4 posterior supplements closer together (Fig. 20E), the others had the posterior 5 set-off. Remarks. Sabatieria similis (Allgtn, 1933) is identical with S. ornata apart from the amphid, which in any case is probably misdrawn in Allgkn s figure-there should almost certainly be more turns than he depicts: the species are therefore synonymized. Sabatieria proabyssah Vitiello & Boucher, also appears identical with S. ornata except for apparent minor details of the gubernaculum structure which are in any case difficult to distinguish: the species are synonymized.

38 64 H. M. PLATT Figure 20. Sabatieria ornafa. A, Amphid; B, cuticle in anterior oesophagus region; C, cuticle in midbody region; D, cuticle in cloaca1 region; E, precloacal supplements and precloacal spine (arrowed); F, 8 posterior region; G, tail; H, head showing R3 sensilla; I, entire 8. Scale bars: A-E, H = 10 pm; F = 50 p; G = 30 prn; I = 300pm.

39 REVISION OF SABATIERIA 65 Records. 14: English Channel (1); North Sea (1); Skagerrak (3); Bresund (4); Norway (1); Mediterranean (4). Allgkn s (1959) two records from the S Atlantic are considered dubious. Sabatieria parabyssalis Wieser, 1954 (Fig. 13; Table 2) Description. Wieser (1954), type. 2 88, Sublittoral fine to coarse sand, m depth, Chilean coast. Remarks. Sabatieria parabyssalis belongs to the subset of the praedatrix group without marked lateral differentiation. Only the amphid and R3 sensilla separates the species from S. ancundiana and S. falczfra. Records. 3: Chile (1): East coast U.S.A. (2). Sabatieria paracupida Wieser & Hopper, 1967 (Fig. 12; Table 2) Description. Wieser & Hopper (1967), type. Cuticle: lateral dots larger and more widely spaced (but not figured). Intertidal sandy-mud, Florida, U.S.A. Remarks. Sabatieria paracupida can only be distinguished by careful comparison of the form of the gubernaculum, amphid size and spicule size. Sabatieria paradoxa Wieser & Hopper, 1967 seems almost identical to S. paracupida apart from having somewhat shorter R3 sensilla (38-46% h.d. versus % h.d.) Records. 1 : Florida, U.S.A. Sabatieria paradoxa Wieser & Hopper, 1967 (Fig. 12; Table 2) Description. Wieser & Hopper ( 1967), type. Intertidal muddy-sand, Florida, U.S.A. Remarks. See previous species. Records. 1 : Florida, U.S.A. Sabatieria pisinna Vitiello, 1970 (Fig. 15; Table 4) Description. Vitiello (1970), type. 3 66, TL = 657 pm. Cuticle without lateral differentiation, but midbody dots smaller than in the cervical and caudal regions. No supplements. Sublittoral mud at 310 m and 410 m depth, French coast, Mediterranean. Remarks. On the basis of the tail shape and form of the gubernaculum, S. pisinna would appear to belong to the pulchra group but is differentiated by the small overall size. Sabatieria pisinna appears to lack the typical paired cervical setae and supplements of the group. But in the same paper, Vitiello (1970) reported another species which normally possesses supplements (viz. S. hizarula) as lacking 5

40 66 H. M. PLATT these organs, so it must remain questionable as to whether S. pisinna was really devoid of precloacal supplements or whether they were overlooked. Records. 2: Mediterranean. Sabatieria praedatrix De Man, 1907 b (Figs 8G, 11D, 12, 21; Table 2) Sabatieria cobbi Kreis, Syn. nov. Sabatieria rugosa Stekhoven, Syn. nov. Material. 1 8 from Exe estuary, SW England: BMNH Descriplions. (1) De Man (1907a), type. 1 8, 1 9: TL = 2900, 3100 pm; a = 50-55, Amphid 2 turns. Anteriorly directed process at distal end of spicules. No supplements observed. Dutch coast. (2) Stekhoven (1946). 1 b, About 13 tiny supplements detected. Skagerrak, Baltic. (2) Additional observations. TL = 1760 pm; a = 38. Cuticle annulated and ornamented with transverse rows of dots: fewer rows of larger dots laterally (Fig. 2 1 C) which may appear longitudinally elongated (Fig. 2 1 D). Cuticularized ventral structure observed at distal end of spicules (arrow in Fig. 21G). Gubernaculum strongly cuticularized (Fig. 2 1 G). Seventeen very small supplements (Fig. 21H), probably easily overlooked. Remarks. Sabatieria praedatrix is chosen as typical of one of the Sabatieria subgroups. Kreis (1929) described a single male from the northwest coast of France (S. cobbi), which had a gubernaculum without apophyses or supplements: otherwise it was very similar to S. praedatrix. Sabatieria praedatrix as described by De Man (1907a) and as observed here possesses well-developed rectal glands which tend to obscure the apophyses. Despite Kreis (1929) generally poor figures, he did indicate glandular structures in the cloaca1 area which may well have obscured the apophyses. On this basis, S. cobbi is considered synonymous with S. praedatrix. Sabalieria rugosa Stekhoven, 1950 from Villefranche only appears to differ from S. praedatrix in having a more filiform tail-otherwise it is indistinguishable and so I propose that they be considered synonymous. Records. 13: North Sea, Skagerrak and Norwegian Coast (7); Kattegat and environs (2); English Channel (1: this paper); Mediterranean (7); Brazil (2). Allgin s (1959) record from the Southern Ocean is considered dubious. Sabatieria Propisinnu Vitiello, 1976a (Fig. 15; Table 4) Description. Vitiello (1976a), type 4 88, Cuticle: lateral differentiation of larger more irregular dots in cervical and caudal regions, smaller in midbody region. Amphid difficult to distinguish. At least 3 supplements observed, but total number not determined. Sublittoral mud, French Mediterranean coast. Remarks. This species belongs to the pulchra group but is rather inadequately

41 REVISION OF SABST1ERI Figure 21. Subutieriu pruedutrzx. A, Amphid; B, cuticle in anterior oesophagus region; C, cuticle in midbody region; D, cuticle in midbody region with elongated punctations; E, cuticle in cloaca1 region; F, entire 6; G, copulatory apparatus showing strongly cuticularized gubernaculum and additional ventral structure (arrow); H, precloacal supplements (arrowed). Scale bars: A-E, G, H = 10 pm; F = 300 pm.

42 68 H. M. PLATT

43 REVISION OF SABATZERZA 69 known. The body size is similar to S. pisinna Vitiello, 1970, which apparently lacks lateral differentiation and has shorter R3 sensilla. Records. 1 : Mediterranean. Sabatieria pulchra (G. Schneider, 1906) (Figs 6C, D, 8C-E, 15, 22; Table 4) Parasabatieria clavicauda Filipjev, syn. Gerlach ( 1965) Parasabatieria vulgaris De Man, 1907b syn. Riemann (1970) Sahalieria quadripapillata sensu Gerlach, 1957b nec Filipjev, Syn. nov. Sabatieria americana sensu Wieser, 1959 nec Timm, Syn. nov. Sabatieria lrivialis Tchesunov, Syn. nov. nec Sabatieria brevisela Stekhoven, 1935a nec Parasabatieria punctata Kreis, 1924 Malerial. 5 88, 5 from Krogarviken, Finland (type locality: leg. P. Jensen, lo.xi.1982), soft bottom, 2 m depth, BMNH , 2 99 from Exe estuary, SW England (leg. R. M. Warwick), BMNH Descriptions. Several descriptions have been made of this species (see Gerlach & Riemann, 1973), mostly under the names S. vulgaris and S. clavicauda. The most recent description is that by Jensen (1979b). Additional observations: anterior gonad left, posterior right of gut. Remarks. Sabatieria pulchra appears well characterized by the cuticle form (Fig. 22A-C), amphid and R3 sensilla size (Fig. 22D-E) and form of the copulatory apparatus (Fig. 22F-G). The various descriptions of S. clavicauda and S. vulgaris appear virtually indistinguishable both from each other and from the albeit rather poor original description of S. pulchra: Jensen s ( ) opinion that S. clavicauda and S. vulgaris constitute valid species cannot be supported on the available morphological evidence. Both S. punctata Kreis, 1924 (synonymized with S. clavicauda by Wieser, 1954) and S. breviseta Stekhoven, 1935a (synonymized with S. pulchra by Lorenzen, 1974) can now be recognized as separate species, as recently described (Platt, 1984). Practical separation is based mainly on the number of amphid turns (3) and cuticle ornamentation. In general, however, the various reports and descriptions of specimens under the names S. pulchra, S. clavicauda, S. vulgaris, S. breviseta and S. punctata are so hopelessly confused that it will never be possible to separate them on any objective basis: it is probably best to disregard most of them. For example, Gerlach s (195713) specimens from Brazil, described as S. clavicauda, would not be S. pulchra as now understood: the lateral cuticle punctations were described as distinct& larger than the median ones. The description probably fits S. punctata Figure 22. Sabatieria pulchra from Krogarviken, Finland. A, Cuticle in anterior oesophagus region; B, ruticle in midbody region; C, cuticle in cloacal region; D, head showing buccal cavity and cephalic sensilla; E, head showing amphid; F, cloacal region showing gubernaculum apophysis, precloacal spine and first supplement; G, copulatory apparatus showing gubernaculum medium piece; H, entire d; I, posterior region showing supplements; J, 0 tail. Scale bars: A-G = 10 pm; H = 300 pm; I, J = 30 pm.

44 70 H. M. PLATT more closely but to transfer it (and others likewise) would be simply an exercise in taxonomic book-keeping. Nevertheless, in a few instances, some reorganization does appear justified. Wieser s (1959) specimens from the west coast U.S.A., described under the name S. americana Timm, 1952, appear to agree with De Man s (1907a) original description of S. vulgaris ( = S. pulchra) : the record is transferred accordingly. Likewise, Gerlach s ( 1957) description of S. quadripapillata Filipjev, 1922 and Tchesunov s (1978) S. trivialis from the Caspian Sea both agree with S. pulchra and are also transferred. Sabatieria punctata (Kreis, 1924) (Figs 6G, 15; Table 4) Sabatieria americana Timm, Syn. nov. Material. 2 d6, 1 from Swansea Bay, Wales: BMNH , d6 from NE England: BMNH Descriptions. (1) Kreis (1924), type. 1 6: TL = 1648 pm; a = 53. (2) Stekhoven (1935a). 2 $6, 2 99: TL = 1520 pm; a = 38. (3) Jensen (1979a) , 37 99: TL = pm. (4) Platt (1984). 4 bd, 1 9: TI. = pm. Testes: anterior right, posterior left of gut. Ovaries: anterior right, posterior left of gut. Remarks. The description by Kreis (1924) is rather poor and could be anything; Stekhoven s (1935a) likewise. However, Jensen (1979a) in his redescription of specimens from the Bresund, Denmark indicated that the species can be separated from S. pulchra mainly by having a larger amphid with more turns and better developed lateral differentiation. Timm s (1952) specimens from Maryland, U.S.A. described as S. americana are, according to the description, indistinguishable from S. punctata and the species are therefore synonymized. Tietjen s (197 1) record of S. americana from the east coast of the U.S.A. is also transferred to S. punctata. Records. 2 1: North Sea, Norwegian coast (8); Baltic, (aresund, Kattegat, Kiel Bay (10); Bristol Channel (1: Platt, 1984); East coast, U.S.A. (2). Sabatieria slekhoueni Vitiello, 1970 (Fig. 12; Table 2) Descriptions. (1) Stekhoven (1950), type. 1 $, 19, TL 6 = 1800 pm; a = 42. Cuticle with larger and more widely spaced dots. Text states 8 amphid with 3$ turns (figure depicts 24); 0 amphid 24 turns. No supplements observed. (2) Vitiello (1970). 3 38, TL = pm; a = Lateral cuticle punctations larger and less numerous than submedian; this differentiation most marked in oesophagus and tail regions and in midbody lateral dots are very small. Amphid with 3-34 turns. Number of supplements difficult to distinguish but apparently Remarks. The tails of the specimens described by Vitiello (1970) have a shorter filiform section than those originally described (as S. abyssalis) by Stekhoven (1950): otherwise, they are similar (assuming that Stekhoven s drawing of the male amphid is correct!). Records 2: Mediterranean.

45 REVISION OF SABATIERIA Sabatieria strigosa Lorenzen, 1972 (Fig. 16; Table 5) Description. Lorenzen (1972), type Sublittoral sand, near Helgoland. Remarks. This species belongs to the celtica group: the curved gubernaculum appears to distinguish it from members of the arrnata group. Records 1: North Sea. 71 Sabatieria supplicans Gerlach, 1956 (Fig. 14; Table 3) Description. Gerlach (1956), type Text states that in the anterior part of the body there are irregular lateral rows of larger punctations. Intertidal sand, Brazil. Remarks. This species appears to belong to the armata group, distinguished from the other species primarily on the size of the copulatory apparatus and tail length. However, information on the number of supplements is wanting. Records. 1: Brazil. Sabatieria triplex Wieser, 1954 (Fig. 12; Table 2) Description. Wieser (1954), type. 1 6, 1 0, 2 juveniles. Cuticle punctations in transverse rows, conspicuously larger laterally. Amphid in male with 4 turns. Sublittoral sediment, 18 m and 200 m depth, Chilean coast. Remarks. This species appears to be very similar to S. praedatrix in cuticle pattern, the copulatory apparatus structure, disposition of supplements and tail shape but has an amphid with one more turn. Records. 3: Chile (1); NW Atlantic (2: Tietjen, 1971, 1976). Sabatieria vasicola Vitiello, 1970 (Fig. 12; Table 2) Description. Vitiello (1970), type. 2 66, Cuticle: lateral differentiation begins about 2 h.d. posterior to the amphid and consists of larger dots which are not arranged in such regular transverse rows as medially. Presence of supplements difficult to distinguish: 19 very small setiform supplements. Sublittoral mud, m depth, French Mediterranean coast. Remarks. The description only provides a subventral view of the male copulatory apparatus and the supplements are described as setiform which is unusual for Sabatieria species. Clearly, more details are needed but based on the tail shape and head structure the species should be recognizable again, especially if collected from near the type locality. Sabatieria uasicola appears to belong to the praedatrix group. Records. 2: Mediterranean.

46 _...I 12 H. M. PLATT , :: * P 9 5 P, clj

47 REVISION OF SABATIERIA 7 :3 Species inquirendae The following eight species cannot unequivocally be allocated to any of the Sabatieria groups through being inadequately described. However, it is not impossible that if collections were made in or near the type localities, then they might be recognized and their position could then be reassessed. In the meantime, they are here considered species inquirendae. The description of each species is briefly reviewed below, data is given in Table 7 and they are illustrated in Fig. 23. The number of specimens used in the original description is given in each case. Sabatieria aspera Sergeeva, 1972: 1 3. Part of cuticle has the appearance of spines (translation from Russian): it is unclear to what this refers. The figure is poor. Location: 85 m depth, mud, Black Sea. Only record. Sabatieria efilata Stekhoven, 1950: 1 6. Poor figure. No supplements observed. If it were not for the slender body, the species could be synonymized with any of several other species depending in the supplement number. Location: Mediterranean. Other records: Tietjen (1969, 1977; east coast U.S.A.). Sabatieria possjetica Platonova, 1971: 10 88, No supplements. Poor figure. Apart from absence (?) of supplements, species indistinguishable from members of the pulchra group. Location: Sea of Japan. Only record. Sabatieria praebosporica Sergeeva, 1972: 3 8.6, 1 9. Cuticle punctated, part of which has the appearance of spines : as with S. aspera, it is unclear what is meant. Poor figure, supplements not figured. Location: 70 m depth, Black Sea. Only record. Sabatieria quadripapillata Filipjev, 1922: 16, 1 9. Cuticle very finely punctated but not figured. Number and disposition of supplements appears characteristic, but description is poor. Location: Sea of AZOV, U.S.S.R. Only valid record. Gerlach s ( 1957) re-description of the species reported 6-7 supplements decreasing in distance apart anteriorly which fits the description of S. pulchra and is therefore transferred accordingly. Sabatieria rota Gerlach, 1957a: 1 6. Cuticle irregularly punctated (not figured). Posterior oesophageal bulb double and amphid loop-shaped: characters unique for Sabatieria although supplement disposition suggests a relationship with pulchra group. Until more information is available, I propose to treat S. rota as a species inguirenda. Location: intertidal sand, Brazil. Only record. Table 7. Differentiating data of species inquirendae (males) Spccies TL a A% R3 Spic Ps T S. aspera S. efiiata.s. possjetica S. praebosporica S. quadripapillata S. rota S. sarcina S. wieseri Abbreviations: see Table ? ?

48 74 H. M. PLATT Subatieriu surcina Vitiello, 1976a: 1 d. Cuticle with larger, more widely spaced punctations laterally. Supplements not observed. Jensen ( 1979a) suggested Vitiello s specimen had a broken tail and this, together with the lack of knowledge on the supplements, suggests more information is required. Location: subtidal sand, Mediterranean. Only record. Sabutieria wieseri nom. nov. for Sabatieria heterura sensu Wieser, 1954 nec Cobb, 1898: 4 dd, Wieser (1954) suggested his specimens were identical with Cobb s (1898) S. heteruru. However, Cobb s single male was not illustrated and little morphometric data were provided. There is insufficient detail in Cobb s description for Wieser (or anyone) to substantiate an identification with S. heteruru: it seems safer to consider Cobb s species as dubious. Wieser s (1954) specimens are therefore considered a distinct species, probably new to science, but inadequately known. Location: Chile. Only record. (See note added in proof.) REFERENCES ALLGEN, C., Uber einige antarktische freilebende marine Nematoden. zoologica Anzeigcr, I: ALLGEN, C., Freilebende marine Nematoden aus dem Drobakabschnittes des Oslofjords. <oologischc Jahrbiicher (Systematik), 61: ALLGEN, C., Freilebende Nematoden aus dem Trondhjemsfjord. Capita <oologica, 4 (2): ALLGEN, C., Zur Kenntnis norwegischer Nematoden 111. Weitere Nematoden aus Tarva. Kongelige Norske Videnrkabernes Selskabs Forhandlinger, 7: ALLGEN, C., Die freilebenden Nematoden des Oresunds. Capita zoologica, 6 (3): ALLGEN, C., Papers from Dr. Th. Mortensen s Pacific Expedition LXXV. West American marine nematodes. Vidmkabelige Meddelelser fra Dansk Naturhistorisk Forening i Xjebenhaun, 110: ALLGEN, C. A., Papers from Dr. Th. Mortensen s Pacific Expedition LXXVI. Pacific freeliving marine nematodes. Videnskabelige Meddelelscr fra Dansk Naturhistorisk Forening i Kjobenhaun, 113: ALLGEN, C., uber eine bemerkenswerte neue Siidsee-Art der Nematodengattung Sabaticria De Rouville, S. heterospculum von Sud-Georgien. Kongelige Norske Vidmkabmes Selskabs Forhandlingcr, 26: 4-6. ALLGEN, C., Freeliving marine nematodes from East Greenland and Jan Mayer. The Swedish Greenland Expedition Meddelelscr om Grenland, 107 (6): ALLGEN, C., Freeliving marine nematoda. Further zoological results of the Swedish Antarctic Expedition ,5 (2): ARNOLD, E. N., Estimating phylogenies at low taxonomic levels. zeifschr$t fur <oofogische qstematik wtd Evolutionsfrschung, 19 (1): BASTIAN, H. C., Monograph on the Anguillulidae, or free Nematoids, marine, land, and freshwater; with descriptions of 100 new species. Transactions of the Linncan Society of London, 25: BOUCHER, G., Distribution quantitative et qualitative des Nimatodes d une station de vase temgene cbtitre de Banyulus-sur-Mer. Cahiers de Biologie Marine, 13: BOUCHER, G., PremiZres donntes tcologiques sur les ntmatodes libres marine d une station de vase cbtitre de Banyuls. Vie Milieu, 23: BRESSLAU, E. & STEKHOVEN, J. H. S., Marine freilebmde Nematoden aus der Nordrcc. Musk royal d Histoire naturelle de Belgique. Bruxelles: COBB, N. A., Australian free-living marine nematodes. Proceedings of the Linncan Society of New South Wales, (2) 23: COBB, N. A., Antarctic marine free-living nematodes of the Shackleton Expedition. Contributions lo a Science of Nernatology, Baltimore, I: COBB, N. A., Marine free living nemas. Australasian Antarctic Expedition , (C) 6 (7): DE CONINCK, L. A., In P.-P. Grasse (Ed.), Classe des Ntmatodes-Systtmatique des Nematodes et sous-classe des Adenophorea. Trait6 de zoologic, 4 (2): DE CONINCK, L. A. & STEKHOVEN, J. H. S., The freeliving marine nemas of the Belgian coast. 11. Mimoires du Mule Royal &Histoire Naturelle de Belgique, 58: DITLEVSEN, H., Marine freeliving nematodes from Danish waters. Videnskabelige Meddcleher fra Dansk Naturhistorisk Forening i Kjebmhavn, 70: DITLEVSEN, H., Papers from Dr. Th. Mortensen s Pacific Expedition Marine free-living nematodes from the Auckland and Campbell Islands. Videnskabelige Meddelelser fra Dansk Naturhistorkk Forening i Kj~benhavn, 73: FILIPJEV, I., Free-living marine nematodes of the Sevastopol area (in Russian). Part I. 74 Osoboi zoologicheskoi Luboratorii i ScuastopoP skoi Biologicheskoi Stantsii, 2 (4):

49 REVISION OF SABATIERZA 75 FILIPJEV, I., Sur les Ntmatodes libres de la mer d Azov Trudy Stauropol skago Sel skokhoqaistuennago Instituta, I: FILIPJEV, I., The classification of the free-living nematodes and their relation to the parasitic nematodes. Smithsonian Miscellaneous Collections, 89 (6): FILIPJEV, I., Ntmatodes libres du bassin polaire (in Russian and French). Dreifuiushchaia ekspeditsiia Glauseumorputi na ledokol nom purokhode G. Sedou Trudy, 3: GERLACH, S. A,, Nematoden aus dem Kiistengrundwasser. Abhandlungen der Mathematisch-. ~aturmissenschaftlichen Klasse. Akademie der Wissenschaften und der Literature Main<, 6: GERLACH, S. A., Die Nematodenbesiedlung des Sandstrandes und des Kustengrundwassers an der italienischen Kuste. I. Systematischer Teil. Archiuio zoologic0 Ztaliano, 37: GERLACH, S. A,, Die Nematodenbesiedlung des tropischen Brandungsstrandes von Pernambuco. Brasilianische Meeres-Nematoden 11. Kieler Meeresjorschungen, 12: GERLACH, S. A,, 1957a. Die Nematodenfauna des Sandstrandes an der Kuste von Mittelbrasilien (Brasilianische Meeres-Nematoden IV). Mitteilungen aus dem zoologischen Museum in Berlin, 33: GERLACH, S. A,, 1957b. Marine Nematoden aus dem Mangrove-Gebiet von Canancia (Brasilianische Meeres -Nematoden 111). Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse. Akademie der Wissenschaften und der Literatur in Main<, 5: GERLACH, S. A., Freilebende Meeresnematoden aus der Gezeitenzone von Spitzbergen. Veroyentlichungen des Instituts fur Meeresjorschung in Bremerhauen, 9: GERLACH, S. A. & REIMANN, F., The Bremerhaven checklist of aquatic nematodes. A catologue of Nematoda Adenophorea excluding the Dorylaimida. Veroffentlichungen des Instituts fur Meeresforschung in Bremerhauen, Supplement 7: pp GOURBAULT, N., NCmatodes abyssaux (Campagne Walda du N/O I. Charcot ). 11. Esptces et genre nouveaux de Comesomatidae. Bulletin du Mudum National PHistoire Naturelle, 2( A), 3: GROZA-ROJANCOVSKI, E., Contributions to the study of free-living nematodes from the Black Sea. Trauaux du Muslum d Histoire Naturelle Gr. Antipa, 13: INGLIS, W. G., Two new species of free-living marine nematodes from the west coast of Scotland. Hydrobiologica, 18: JAYASREE, K. & WARWICK, R. M., Free-living marine nematodes of a polluted sandy beach in the Firth of Clyde, Scotland. Description of seven new species. Journal of Natural History, 11: JENSEN, P., 1979a. Revision of Comesomatidae (Nematoda). zoologica Scripta, 8: JENSEN, P., 1979b. Nematodes from the brackish waters of the Southern archipelago of Finland. Benthic species. Annales zoologici Fennici, 16: JENSEN, P., Species, distribution and a microhabitat theory for marine mud dwelling Comesomatidae (Nematoda) in European waters. Cahiers de Biologie Marine, 22: JENSEN, P. & GERLACH, S. A., Three new Nematoda-Comesomatidae from Bermuda. Ophelia, 16 (1): KREIS, H. A., Zur Kenntnis der freilebenden marinen Nematoden. Schriftn fur Susswasser-und Meereskunde, Busum, 2: KREIS, H. A., Freilebende marine Nematoden von der Nordwestkiiste Frankreichs (Trtbeurden Cdtes du Nord). Capifa zoologica, 2 ( I): LORENZEN, S., Die Nematodenfauna im Verklappungsgebiet fir Industrieabwasser nordwestlich von Helgoland. 11. Desmodorida und Chromadorida. zoologische Anzeiger, (1971) 187: LORENZEN, S., Die Nematodenfauna der sublitoralen Region der Deutschen Bucht, insbesondere im Titan-Abwassergebiet bei Helgoland. Verofentlichungen des Instituts fur Meeresjorschung in Bremerhaum, 14: LORENZEN, S., Entwurfeines phylogenetischen Systems der freilebenden Nematoden. Veroffentlichungen des Instituts fur Meeresjorschung in Bremerhaum, Supplement 7: pp DE MAN, J. G., 1907a. Sur quelques esptces nouvelles ou peu connues des NCmatodes libres habitant les cdtes de la Ztlande. Mlmoires de la Sociltt! <oologique de France, 20: DE MAN, J. G., 1907b. Sur quelques esptces nouvelles ou peu connues des Ntmatodes libres vivant sur les cbtes de la Ztlande. Tijdschrifi der Nederlandsche Dierkundige Vereeniging, 10: DE MAN, J. G., Neue freilebende nematoden aus der Zuidersee. Tijdschriyt der Nederlandsche Dierkundige Vereeniging, (2) 18: MICOLETZKY, H., Letzter Bericht uber freilebende Nematoden aus Suez. Sitzungsberichte der Akademie der Wissenschaften. Mathematischen-Naturwissenschaftliche Classe. Wien, 133: PLATONOVA, T. A,, Free-living marine nematodes from the Possjet Bay of the Sea of Japan (in Russian). Fauna iflora zaliua Posjeta japnskovo morja: PLATT, H. M., Revision of the Ethmolaimidae (Nematoda: Chromadorida). Bulletin of the British Museum (Natural History) zoology series, 43: PLATT, H. M., The freeliving marine nematode genus Sabatieria (Nematoda: Comesomatidae). I. Two new species from Stonington Island, Antarctica. Bulletin of the British Museum (Natural History) zoology series, 45: PLATT, H. M., The freeliving marine nematode genus Sabatinia (Nematoda: Comesomatidae). 11. Redexriptions of five European species. Bulletin of the British Museum (Natural History) zoology series, 46:

50 76 H. M. PLATT RIEMANN, F., Die interstitielk Fauna im Elbe-Aestuar Verbreitung und Systematik. Archiu fur Hydrobiotogie, Supplement 31: RIEMANN, F., Freilebende Nematoden aus dem Grenzbereich Meer-Siiss-Wasser in Kolumbien, Siidamerika. Verojimtlichungen des Instituts fur Meeresfrchung in Bremerhauen, 12: ROUVILLE, E. de, Enumeration des Nimatodes libres du canal des Bourdignes (Cette). Compte Rmdu des Slances de la Socilti de Biologic, 55: SCHNEIDER, G., Beitrag zur Kenntnis der im Uferschlamm des Finnischen Meerbusens freilebenden Nematoden. Acta Societalis pro Fauna et Flora Fennica, 27 (7): SERGEEVA, N. G., New species of free-living nematodes from the order Chromadorida in the Black Sea (in Russian). <oologicheskii Zhurnal51 (8): SERGEEVA, N. G., The structure of complexes of free-living marine nematodes in a Modioluspheolinus biocoenosis (in Russian). Biolog&a Morya, 36: SOUTHERN, R., Nemathelmia, Kinorhyncha and Chaetognatha (Clare Island Survey, part 54). Proceedings of th Royal Irish Academy, 31: SSAWELJEV, S., Zur Kenntnis der freilebenden Nematoden des Kolafjords und des Relictensee Mogilnoje. Trudy Imperalorskugo Sankt-Peterburgskugo Obschchestua Estestuoispytatelci, 43: STEINER, G., Freilebende Nematoden aus der Barentsee. <oologische Jahrbiichcr (Systemafik), 39: STEKHOVEN, J. H. S., 1935a. Additional notes to my monographs on the freeliving marine nemas of the Belgian Coast. I. and 11. Mlmoires du Musle Royal PHistoire Naturelle de Belgique, 72: STEKHOVEN, J. H. S., 1935b. Nematda: Systamatischer Ted, Nematode errantia. In G. Grimpe & E. Wagler (Eds), Die Tierwelt der Nord-xnd Ostsee (Leipzig 1935), 5b STEKHOVEN, J. H. S., Freilebende marine Nematoden des Mittelmeeres. IV. Freilebende marine Nematoden der Fishchereigriinde bei Alexandrien. <oologische Jarhbiicha (Systematik), 76: STEKHOVEN, J. H. S., Freilebende marine Nematoden des Skageraks und der Umgebung von Stockholm. Arkiu fur <oologi 37A: 1-91, STEKHOVEN, J. H. S., The freeliving marine nemas of the Mediterranean. I. The Bay of Villefranche. Mimoires de l hstitut Royal des Sciences Naturelles de Belgigue, 37: TCHESUNOV, A. V., New free-living nematodes from the Caspian Sea (in Russian). <oologischeskiz zhurnal, 57: TIETJEN, J. H., The ecology of shallow water meiofauna in two New England estuaries. Occologia (Berlin), 2: TIETJEN, J. H., Ecology and distribution of deep-sea meiobenthos off North Carolina. Deep Sea Research, 18: TIETJEN, J. H., Distribution and species diversity of deep-sea nematodes off North Carolina. Deep Sea Research, 23: TIETJEN, J. H., Population distribution and structure of freeliving nematodes of Long Island Sound. Marine Biology, 43: TIETJEN, J. H., Population structure and species composition of the free-living nematodes inhabiting sands of the New York Bight apex. Estuarine and Coastal Marine Science, 10: TIMM, R. W., A survey of the marine nematodes of Chesapeake Bay, Maryland. Contributiomfrom f/ze Chesapeake Biological Laboratory, 95: TIMM, R. W., The marine nematodes of the Bay of Bengal. Proceedings of the Pakistan Academy of ScimC, 1: TIMM, R. W., Some estuarine nematodes from the Sunderbans. Proceedings of the Pakistan Academy of Science, 4: VITIELLO, P., Nimatodes libres marins des vases profondes du Golfe du Lion. 11. Chromadorida. Tlthys, 2: VITIELLO, P., Peuplements de nirnatodes marins des fonds envasis de Provence. I.-Skliments vaseux de mode calme et vases terrigines cbtiires. Annales de l lmtitut Oclanographiqu, 50: VITIELLO, P., 1976a. Trois nouvelles espices de Comesomatidae (Nematoda) des CBtes de Provence. Tethys, 7: VITIELLO, P., 1976b. Peuplements de nimatodes marins des fonds envasis de provence. 11. Fonds ditritiques envasis et vases bathylaes. Annales de I lnstitut Oclanographique, 52: I. VITIELLO, P. & BOUCHER, G., Nouvelles esptces de Chromadorida (Nematoda) des vases terrigines Mediterraniennes. Bulletin de la Sociltl <oologigue de France, 96: WARD, A. R., Three new species of free-living marine nematodes from sublittoral sediments in Liverpool Bay. Marine Biology, 24: WARWICK, R. M., Freeliving marine nematodes from the Indian Ocean. Bulletin of the British Museum (Natural History) oology series, 25: WARWICK, R. M., Some free-living marine nematodes from the Isles of Scilly. Journal ojnatura1 ~ist~ry, 11: WARWICK, R. M. & COLES, J. W., The marine flora and fauna of the Isles of Scilly. Free-living Nematoda. Journal of Natural History, 11: WIESER, W., Free-living marine nematodes. 11. Chromadoroidea. Acta Uniuersitatis Lundmis, N.F.Z., 50:

51 REVISION OF SABATIERZA 77 WIESER, W., Free-living Nematodes and other Small Invertebrates of Puget Sound Beaches. Seattle: University of Washington Press. WIESER, W. & HOPPER, B., Marine nematodes of the east coast of North America. 1. Florida. Bulletin of the Museum of Comparatiue ~oologv, Haruard College, 135: APPENDIX List of ualid Sabatieria species S. abyssalis (Filipjev, 1918) S. alata Warwick, 1973 S. ancudiann Wieser, 1954 S. armata Gerlach, 1952 S. brevisefa Stekhoven, 1935a S. celtica Southern, 1914 S. conicauda Vitiello, 1970 S. demani Bresslau & Stekhoven in Stekhoven, 1935b S. dodecaspapillata (Kreis, 1929) S. elongafa Jayasree & Warwick, 1977 S. falcifera Wieser, 1954 S. furcillata Wieser, 1954 S. granger Wieser, 1954 S. infermissa Wieser, 1954 S. kefletti Platt, 1983 S. lawsi Platt, 1983 S. longisetosa (Kreis, 1929) S. longispinosa Lorenzen, 1972 S. lyonessa Warwick, 1977 S. macramphis Lorenzen, 1972 S. migrans Jensen & Gerlach, 1977 S. mortenreni (Ditlevsen, 1921) S. ornata (Ditlevsen, 1918) S. parabyssalis Wieser, 1954 S. paracupida Wieser & Hopper, 1967 S. paradoxa Wieser & Hopper, 1967 S. pirinna Vitiello, 1970 S. praedafrix De Man, 1907b S. propisinna Vitiello, 1976 S. pulchra (G. Schneider, 1906) S. punctata (Kreis, 1924) S. sfekhoueni Vitiello, 1970 S. strigosa Lorenzen, 1972 S. supplicans Gerlach, 1956 S. triplex Wieser, 1954 S. vasicola Vitiello, 1970 List of species inquirendae S. aspera Sergeeva, 1972 S. ejilafa Stekhoven, 1950 S. possjefica Platonova, 1971 S. praebosporica Sergeeva, 1972 S. quadripapillata Filipjev, 1922 S. rofa Gerlach, 1957a S. sarcina Vitiello, 1976a S. luieseri nom. nov. List of dubious species S. antarctica Cobb, 1914 juveniles only described; considered doubtful by Wiescr (1954). S. arcfica (Allgkn, 1954): poor figure Dub. nov. S. arcuafa Wieser, 1954: based on single damaged specimen. Dub. nov. S. ausfralis AllgCn, 1929: based on juveniles only; considered dubious by Platt (1983). S. cettensis Rouville, 1903: considered doubtful by Filipjev (1922) and Wieser (1954). S. cleopafris Micoletzky, 1924: 99 only, no figure. Dub. nov. S. Jilicaudafa Allgtn, 1951: considered doubtful by Wieser (1954). S. heterospiculum Allgtn, 1953: considered dubious by Platt (1983). S. heferura (Cobb, 1898) nec Wieser 1954, no illustration. Dub. POV. S. jubufa Cobb, 1898: insufficient description. Dub. nov. S. kolacnsis (Ssaweljev, 1912): no illustration. Dub. POV. S. mawsoni Wieser, 1954 pro S. antarctica (Cobb, 1930): poor figure. Dub. UOV. S. norwegica Allgtn, 1931: considered incertae sedis by Wieser (1954) Dub. nov. S. parauufgari.r Filipjev, 1946: Q only. Dub. nov. S. pellucida AllgCn, 1959: 0 only. Dub. nov. S. rotundicauda AllgCn, only. Dub. nov. S. faboguillensis (Allgtn, 1947): considered insufficiently described by Wieser (I 954). Dub. nov. 3. tenuicaudofa (Bastian, 1865): insufficient description; considered dubious by U ieser ( 1954). S. fenuiseta AllgCn, 1959: 9 only; considered inquirenda by Jensen (1979a). Dub. nov. S. fubilaima Allgkn, 1959: 0 only; considered inquirenda by Jensen (1979a). Dub. nov.

52 78 H. M. PLATT List of Setoaabatietia gen. nov. Sabatieria americana Timm, 1952 Sabatieria americana sensu Wieser, 1959 Sabatieria chitwoodi Wieser, 1954 Sabafieria cobbi Kreis, 1929 Sabatieria jibulata Wieser, 1954 Sabatieria granulosa Vitiello & Boucher, Sabatieria heterura smu Wieser, 1954 Sabatieria hilarula De Man, 1922 Sabafieria jubata sensu Wieser, 1954 Sabatieria microseta Timm, 1967 Sabatieria proabssalis Vitiello & Boucher, Sabatieria quadripapillafa semu Gerlach, 1957 Sabatieria rugosa Stekhoven, 1950 Sabatieria scotlandia Inglis, 1961 Sabatieria similis (Allgtn, 1933) Sabafieria triuialis Tchesunov, 1978 new synonyms = Sabafieria punctata (Kreis, 1924). Syn. nov. = Sabatieria pulchra (C. Schneider, 1906). Syn. nov. = Sefosabatieria hilarula (De Man, 1922). Syn. nov. = Sabafieria praedatrix De Man, Syn. nov. = Setosabatieriafibulata (Wieser, 1954). Syn. nov. = Sabatieria granifcr Wieser, Syn. nov. = Sabatieria wieseri. Syn. nov. = Setosabatieria hilarula (De Man, 1922). Syn. nov. = Setosabatieria hilarula (De Man, 1922). Syn. nov. = Pierrickia microseta (Timm, 1967). Syn. nov. = Sabatieria omla (Ditlevsen, 1918). Syn. nov. = Sabafieria pulchra (C. Schneider, 1906). Syn. nov. = Sabatieria praedafrix De Man, Syn. nov. = Sefosabatieria hilarula (De Man, 1922) Syn. nov. = Sabatieria ornata (Ditlevsen, 1918). Syn. nov. = Sabatieria pulchra (C. Schneider, 1906). Syn. nov. NOTE ADDED IN PROOF Whilst this paper was in press, Pastor de Ward (1984) published descriptions of several marine nematodes from Argentina, including three species of Sabatieria. For the sake of completeness, they are listed below with a few brief comments. Sabatieria mortenseni (Ditlevsen, 192 1) Remarks. These relatively large specimens (TL = pm) belong to the pulchra group. Points of difference between Pastor de Ward s specimens and Ditlevsen s original description would appear to be relative length of the R3 sensilla (50% versus 80%) and the conspicuous cuticle punctations present in the Argentinian form. Sabatieria heterura (Cobb, 1898) Remarks. As argued above, I consider S. heterura of Cobb (1898) as dubious. The specimens described by Wieser (1954) from Chile were renamed S. wieseri. Pastor de Ward s specimens would appear to be similar to those of Wieser (1954) and it would seem reasonable to consider them conspecific. Obviously in view of Pastor de Ward s excellent description, S. wieseri cannot now be considered species inquirenda and should be incorporated into the praedatrix group. As pointed out by Pastor de Ward, the Argentinian specimens are also very similar to S. lawsi described by Platt (1983) from Antarctica. Further examination of the specimens might result in their being synonymized. Sabatieria sp. Remarks. These specimens belong to the celtica group but would appear to have points of difference from all known valid species. REFERENCES PASTOR DE WARD, C. T., Nematodes marinos de la Ria Deseado (Axonolaimoidea: Axonolaimidae, Diplopeltidae, Comesomatidae) Santa Cruz, Argentina. 4. Centro Nacionat Patagdnico Conseio Nacionaf de Investigaciones CientiJicas y TCcnicas, Contribucion Nro. 86: pp. 2 1.